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AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
~ WYCOTAXON
Volume L January-March 1994
CONTENTS
Contribution to our knowledge of the Aphyllophorales (Basidiomycotina) of the Basque
MOIIREN ee ge a aig iwi Where ata a se sckeieo arelia ei I, Salcedo Larralde 1
Contribution to the lichen flora of Transkei........... C. M. Thomas and R. B. Bhat 9
Battarreoides diguetti (Gasteromycetes, Tulostomatales) in Southern Africa.
J. C. Coetzee and A. Eicker 19
Type studies in the Polyporaceae 25. Species described from Japan by R. Imazeki and A. Yasuda.
Tsutomu Hattori and Leif Ryvarden 27
The species of the genus Rinodina (Lichenized Ascomycetes, Physciaceae) containing
Pannarin in Eurasia with a special note on the taxonomy of Rinodina granulans.
Mireia Giralt, Helmut Mayrhofer and Walter Obermayer 47
Systematic and biological studies in the Balansieae and related anamorphs. VIII.
TheEphelis anamorph of Balansia epichloe.
Ryan A. Phelps and Gareth Morgan-Jones_ 61
A new species of Ascotricha with non-ostiolate ascomata.
Shun-ichi Udagawa, Shigeru Uchiyama and Seigo Kamiya 73
Talaromyces spectabilis, a new species of food-borne ascomycetes.
Shun-ichi Udagawa and Shoji Suzuki 81
Notes on the genus Monoicomyces (Laboulbeniales, Ascomycotina): On the species
Meee IDCO Yr ODGRAZZIM ye ee eo Ba eg, Sergio Santamaria 89
Observations on Pseudotracylla species.......... 0... ce ee L. M. Carris 93
Acaulospora excavata sp. nov. - an endomycorrhizal fungus from Céte d'Ivoire.
K. Ingleby, C. Walker and P. A. Mason 99
Studies on keratinophilic fungi. Il. Chrysosporium pilosum sp. nov.
J. Gené, J. Guarro, K. Ulfig, P. Vidal and J. Cano 107
A new species of Delitschia from West Africa.
Fernando Pelaez, Jon D. Polishook, Marti Valldosera and Josep Guarro 115
A new species of Maravalia from India... .G. Bagyanarayana and E. John Ravinder 123
A new species of Uromyces on Ocimum from India
G. Bagyanarayana and E. John Ravinder 127
Amanita westii - taxonomy and distribution. A rare species from states bordering
the Gulf of Mexico. .............. Rodham E. Tulloss and David P. Lewis 131
Orpinomyces intercalaris, anew species of polycentric anaerobic rumen fungus from cattle.
Y. W. Ho, N. Abdullah and S. Jalaludin 139
The original collections of Arturo Nannizzi (1877-1961) in the Herbarium Universitatis
DeMensis (HIE NAY re ee A. Chiarucci and M. G. Mariotti 151
Lambertellinia scutuloides (Sclerotiniaceae), a new genus and species for a discomycete
previously confused with Hymenoscyphus caudatus.
Richard P. Korf and Pavel Lizon 167
[Contents continued overleaf]
ISSN 0093-4666 MY XNAB 50s 1-516° (1994)
Published quarterly by MYCOTAXON, LTD., P. O. Box 264, Ithaca, Ny 14851.
For subscription details, availability in microfilm and microfiche,
and availability of articles as tear sheets, see back cover.
[Contents continued from front cover]
Agaricales of Baja California - Mexico.
M. Candusso, A. Gennari and N. Ayala 175
Phaulomyces simplocariae sp. nov. (Ascomycetes, Laboulbeniales) from Simplocaria
semistriazd (Coteoiera, Pytrinidges 24 fs i's Be ek oe eR ee A. De Kesel 191
Stomiopeltis glochidiicola sp.nov..............+.+-. Alaka Pande and V. G. Rao 199
Additional new species and new reports of Pertusaria (Lichenised Ascomycotina) from New
Zealand with a revised key to the corticolous species in New Zealand.
Alan W. Archer and John A. Elix 203
Alternaria themes and variations (74-105) .................. Emory G. Simmons 219
Agaricus pseudoargentinus n. sp. from Argentina.
Edgardo Albert6 and Jorge E. Wright 271
Three new Southamerican species of Bovista (Gasteromycetes).
V. L. Suarez and J. E. Wright 279
Four new species in the lichen genus Parmelia (Ascomycotina, Lecanorales) from Southern
Africa, with notes on Southern African lichens.............. Franklin A. Brusse 291
Furia shandongensis (Zygomycetes: Entomophthorales), a new pathogen of earwigs.
Weiman Wang, Wenhua Lu and Zengzhi Li 301
Two new pathogens of dipteran insects ............. Meizhen Fan and Zengzhi Li 309
A new Cortinarius from a mature aspen stand in Montana.
Cathy Cripps and Orson K. Miller, Jr. 315
The lichen flora of Rock Canyon, Utah County, Utah.
Ted O. Ririe, Larry L. St. Clair and Clayton C. Newberry 323
Validation of the name Microbotryum vinosum (Ustilaginales)...Cvetomir M. Denchey 331
New species of Gymnopilus (Agaricales, Cortinariaceae) from Mexico.
Laura Guzman-Davalos 333
Discosia’ eucalypiicola. wan -sponOw ose 8 se a ee T. R. Nag Raj 349
Tricholosporum in Mexico and description of a new species.
Gaston Guzman, Victor M. Bandala and Leticia Montoya 355
Studies in the genus Pleurotus. Ill. The varieties of P. ostreatus -complex based
on interbreeding strains and the study of basidiomata obtained in culture.
Gast6n Guzman, Leticia Montoya, Gerardo Mata and Dulce Salmones 365
Omphalina sensu lato in North America. 1-2. 1.O0mphalina wynniae and the genus
Chrysomphalina. 2.Omphalina sensu Bigelow.
Lorelei L. Norvell, Scott A. Redhead, Joseph F. Ammirati 379
Alternaria themes and variations (106-111) ................. Emory G. Simmons 409
Menispora convoluta, a new dematiaceous hyphomycete from Central Italy.
Dario Lunghini 429
Redisposals and redescriptions in the Monochaetia-Seiridium, Pestaltia-Pestalotiopsis
complexes. X. Pestalotia granati and Pestalozzina punicae
T. R. Nag Raj and V. Mel'Nik 435
Cylindrocladium naviculatum sp. nov. and two new vesiculate hyphomycete genera,
Falcocladium and Vesicladiella.
P. W. Crous, M. J. Wingfield, A. C. Alfenas and S. F. Silveira 441
Wentiomyces lichenicola subsp. nov.bouteillei champignon lichénicole non lichénisé
(Dothideales, Dimeriaceae).
Claude Roux, Olivier Bricaud, Emmanuél Sérusiaux et Clother Coste 459
Miscellaneous notes on Mucoraceae........... M.A.A.Schipper and R.A.Samson 475
hawrnctond ig Anthote. 5 6 Sante on alslt lot vd wiege sl eae koe ot A 2 ee ee 493
Aiitiay TINE eo eG Bits Ga cig lh ett Se oa et ERY eile Wik Ce melee ie a 499
SEES: 10 fi ae AR rail dee atin ae) ee! atte wl nice WW) Nt elle ta fieeh dy Al ens le 502
[og se Ur pelt on 2 Ae ean ds 4 AU Ye Meg” ee Ce Nee MENDON a PIES $15
Publication Date; MY COTAAON Volume 49. 206.0025 os oo Oe eo ee eee 515
Rees Oe ea ee ie Re as Ge seta Uh aoe oe eRe Le gone ee er 516
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AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
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111
Table of Contents, Volume Fifty
January-March 1994
Contribution to our knowledge of the Aphyllophorales (Basidiomycotina) of the Basque
Countrysa lle 3 oe tee Me Eee Pee ee I. Salcedo Larralde 1
Contribution to the lichen flora of Transkei. ....C. M. Thomas and R. B. Bhat 9
Battarreoides diguetti (Gasteromycetes, Tulostomatales) in Southern Africa.
J. C. Coetzee and A. Eicker 19
Type studies in the Polyporaceae 25. Species described from Japan by R. Imazeki
aNd ACRY ASNdae qs cmeah ae ton pode Tsutomu Hattori and Leif Ryvarden 27
The species of the genus Rinodina (Lichenized Ascomycetes, Physciaceae) containing
Pannarin in Eurasia with a special note on the taxonomy of Rinodina granulans.
Mireia Giralt, Helmut Mayrhofer and Walter Obermayer 47
Systematic and biological studies in the Balansieae and related anamorphs. VIII.
TheEphelis anamorph of Balansia epichloe.
Ryan A. Phelps and Gareth Morgan-Jones_ 61
A new species of Ascotricha with non-ostiolate ascomata.
Shun-ichi Udagawa, Shigeru Uchiyama and Seigo Kamiya 73
Talaromyces spectabilis, a new species of food-borne ascomycetes.
Shun-ichi Udagawa and Shoji Suzuki 81
Notes on the genus Monoicomyces (Laboulbeniales, Ascomycotina): On the species
cdlescnbediby SDePaZZiNi wie ek ea eet © eee Sergio Santamaria 89
Observations on PS€udoIacylla ispecies \ rast tar Gowint ona) ales L. M. Carris 93
Acaulospora excavata sp. nov. - an endomycorrhizal fungus from Céte d'Ivoire.
K. Ingleby, C. Walker and P. A. Mason 99
Studies on keratinophilic fungi. Il. Chrysosporium pilosum sp. nov.
Gené, J. Guarro, K. Ulfig, P. Vidal and J. Cano 107
A new species of Delitschia from West Africa.
Fernando Pelaez, Jon D. Polishook, Marti Valldosera and Josep Guarro 115
A new species of Maravalia from India.
G. Bagyanarayana and E. John Ravinder 123
A new species of Uromyces on Ocimum from India :
G. Bagyanarayana and E. John Ravinder 127
Amanita westii - taxonomy and distribution. A rare species from states bordering
the<Gulf'ok MEXICO 3.005... Rodham E. Tulloss and David P. Lewis 131
Orpinomyces intercalaris, a new species of polycentric anaerobic rumen
FONGUS OM Callens ca. oar, ner Y. W. Ho, N. Abdullah and S. Jalaludin 139
The original collections of Arturo Nannizzi (1877-1961) in the Herbarium Universitatis
SETCUSIS COLEINA) b ecw Ret tect eo vei A. Chiarucci and M. G. Mariotti 151
Lambertellinia scutuloides (Sclerotiniaceae), a new genus and species for a
discomycete previously confused with Hymenoscyphus caudatus.
Richard P. Korf and Pavel Lizon 167
Agaricales of Baja California - Mexico.
M. Candusso, A. Gennari and N. Ayala 175
Phaulomyces simplocariae sp. nov. (Ascomycetes, Laboulbeniales) from Simplocaria
somusmiaia (Coleoptera, Byrrhidae) jcc 2. eles elena ees 3+ A. De Kesel 191
Stomiopeltis glochidiicola sp. nov............ Alaka Pande and V. G. Rao 199
Additional new species and new reports of Pertusaria (Lichenised Ascomycotina)
from New Zealand with a revised key to the corticolous species in New Zealand.
Alan W. Archer and John A. Elix 203
Alternaria themes and variations (74-105)............. Emory G. Simmons 219
Agaricus pseudoargentinus n. sp. from Argentina.
Edgardo Alberto and Jorge E. Wright 271
Three new Southamerican species of Bovista (Gasteromycetes).
V. L. Suarez and J. E. Wright 279
iv
Four new species in the lichen genus Parmelia (Ascomycotina, Lecanorales)
from Southern Africa, with notes on Southern African lichens.
Franklin A. Brusse
Furia shandongensis (Zygomycetes: Entomophthorales), a new pathogen of earwigs.
Weiman Wang, Wenhua Lu and Zengzhi Li
Two new pathogens of dipteran insects........ Meizhen Fan and Zengzhi Li
A new Cortinarius from a mature aspen stand in Montana.
Cathy Cripps and Orson K. Miller, Jr.
The lichen flora of Rock Canyon, Utah County, Utah.
Ted O. Ririe, Larry L. St. Clair and Clayton C. Newberry
Validation of the name Microbotryum vinosum (Ustilaginales).
Cvetomir M. Denchev
New species of Gymnopilus (Agaricales, Cortinariaceae) from Mexico.
Laura Guzman-Davalos
Discosia. €ucalypticola "anain-spyNoviniee see cnee eee es T. R. Nag Raj
Tricholosporum in Mexico and description of a new species.
Gaston Guzman, Victor M. Bandala and Leticia Montoya
Studies in the genus Pleurotus. Ill. The varieties of P. ostreatus -complex based
on interbreeding strains and the study of basidiomata obtained in culture.
Gaston Guzman, Leticia Montoya, Gerardo Mata and Dulce Salmones
Omphalina sensu lato in North America. 1-2. 1.Omphalina wynniae and the genus
Chrysomphalina. 2.Omphalina sensu Bigelow.
Lorelei L. Norvell, Scott A. Redhead, Joseph F. Ammirati
Alternaria themes and variations (106-111)............ Emory G. Simmons
Menispora convoluta, a new dematiaceous hyphomycete from Central Italy.
, Dario Lunghini
Redisposals and redescriptions in the Monochaetia-Seiridium, Pestaltia-Pestalotiopsis
complexes. X. Pestalotia granati and Pestalozzina punicae
T. R. Nag Raj and V. Mel'Nik
Cylindrocladium naviculatum sp. nov. and two new vesiculate hyphomycete genera,
Falcocladium and Vesicladiella.
P. W. Crous, M. J. Wingfield, A. C. Alfenas and S. F. Silveira
Wentiomyces lichenicola subsp. nov.bouteillei champignon lichénicole non lichénisé
(Dothideales, Dimeriaceae).
Claude Roux, Olivier Bricaud, Emmanuél Sérusiaux et Clother Coste
Miscellaneous notes on Mucoraceae...... M.A.A.Schipper and R.A.Samson
Instructions tov Aithors.i® Peete ee oe Sas sca eel lee gar ec sti ere acme
Author INDEX). 2952 Sree Nasr ee MA are ae Ha J 1 va eee
INDEX+to Tungous taxdy are eer he reas Nitta porteetaeke «5 2 es NGRa ete ee ee
Errataicys 20s et EO Sel aes MEET UR SEEN S 4, (ig bY a SWE ee
Publication Date,’MYCOPAXON#*Volume 49 25. 03 oka ei
REVICWETS ... 5.551 Re ee OMT le Oth ety tg ae gh RUMORS TL eee Let aed UA Re Rtn ane ne
MY COTAXON
Volume L, pp. 1-7 January-March 1994
CONTRIBUTION TO OUR KNOWLEDGE OF THE
APHYLLOPHORALES (BASIDIOMYCOTINA) OF THE BASQUE
COUNTRY.|II”
|. SALCEDO _LARRALDE
Lab. Botanica. Dpto. Biologia Vegetal y Ecologia. Facultad de Ciencias.
Universidad del Pais Vasco/EHU. Apdo. 644. 48080 Bilbao. Spain.
Summary
Four species until the present unknown in the Iberian mycological flora
are mentioned and described, namely: Acanthobasidium phragmitis Boidin
& al., Auriporia aurulenta A.David & al., Flaviporus brownei (Humb.)
Donk and Sarcodontia crocea (Schwein.:Fr.) Kotl.
Resdmen
Se citan y comentan 4 especies no conocidas hasta ahora para la flora
micolégica ibérica, a saber: Acanthobasidium phragmitis Boidin & al.,
Auriporia aurulenta A.David & al., Flaviporus brownei (Humb.) Donk y
Sarcodontia crocea (Schwein.:Fr.) Kotl.
INTRODUCTION
Following work started a few years ago, which has the final aim of
cataloguing the Basque Country's Aphyllophorales, different phases
corresponding to the sampling of concrete areas have been covered (cf.
Salcedo Larralde, 1989; Salcedo & Telleria, 1992). During the
sampling done, the above mentioned species have turned out to be
interesting, whose appearance might have been favoured by the
climatology of 1992.
The samples studied are in the Herbarium of the University of the Basque
Country (BIO), or in V. Martinez's private Herbarium (VM).
* Work supported by the University of the Basque Country (UPV/EHU) project n°
118.310-E191/91
2
Acanthobasidium phragmitis Boidin & al. Bull. Soc. Mycol. France
101(4):345. (1985)
VIZCAYA: Lejona, barrio deTelleria, 30TWP0098, 50 m, on Arundo
donax, 12-XI-1992, |. Salcedo, 6275IS, BlO-Fungi 4693; idem,
6278IS, BlO-Fungi 4692.
In 1985 Boidin & al. described this species, that forms little
menbranous and whitish patches, on stems of two genera of the Poaceae
family. The specimens studied have also been collected in this same
family and relatively near from the described area.
The microscopic characters are very gaudy, having
pleuroacanthobasidioles with numerous apical protuberances (figs.
1A,3A), also gloeocystidia with a round jutting apex and which are
sometimes rather moniliformic (figs. 1C,3B), about 20-24 x 8-10
um. Basidia have four sterigmata and with many visible protuberances
on the lower middle part, which differentiates it from A. norvegicum
(J.Eriksson & Ryvarden) Boidin & al., since its basidia have two
sterigmata and very few protuberances (l.c.).
The basidiospores, that shrivel up and easily become deformed, are
ellipsoid, very ornamented, and with long spines, easily seen in Melze's
reagent, as they are strongly amyloid, and about 11-13 x 6.5-8 wm
(figs. 1D,3C,3D).
Auriporia aurulenta A.David & al. Bull. Soc. Mycol. France
90(4):359-370 (1974)
ALAVA: Valdegovia, Nograro, 30TVN9040, 950 m, on Pinus sylvestris,
30-V-1992, J.L.Garcia, L.Manso, V.Martinez, X. Olano & |. Salcedo,
7561VM; idem, 5629IS, BlO-Fungi 4690.
GUIPUZCOA: Aya, Laurgain, 30TWN6787, 450 m, on Pseudotsuga
menziesii, 21-X|-1992, J.L. Albizu, J. Huarte, X. Laskibar, J.M.
Lekuona & |. Salcedo, 6336IS, BlO-Fungi 4696.
This species is easily recognized in the field by its basidiocarp
resupinate with hymenophore poroid, deep yellow-orange in colour, but
turning ochre with age.
Its microscopy is distinguished by its monomitic hyphal system, hyphae
with clamps, its cystidia stalks and ventricose, more or less fusiform,
thick-walled, and about 20-35 x 8-12 um, with grainy crystals on top
(figs. 2A,3F) and by the presence of hyaline and lageniformic
leptocystidia, 60-70 x 5-10 um (figs. 2B,3F). The basidiospores are
ellipsoid and, in the specimens studied, about 4.5-6 x 2-3 ym (figs.
CC Slssa)s
10 um
——— et
Fig. 1.- Acanthobasidium phragmitis, BlO-Fungi 4693. A. pleuroacantho-
basidioles. B. basidium. C. gloeocystidium. D. spores.
A
10 pm B
Fig. 2.- Auriporia aurulenta, BlO-Fungi 4690. A. Cystidia with crystals on
top. B. leptocystidium. C. basidium and spores.
4
This species is known in Europa from Austria, Czechoslovakia, France,
and the one time Yugoslavia (cf. Julich 1984:335).
Flaviporus brownei (Humb.) Donk Persoonia 1: 189 (1960)
GUIPUZCOA: Oiartzun, Arizabalo, 100 m, 30TWN9295, on Pinus sp.
very wet, 26-IX-1992, |. Salcedo, 5890IS, BlO-Fungi 3821. Aya,
Laurgain, 30TWN6787, 450 m, on Larix sp., 21-XI-1992, J.L.
Albizu, J. Huarte, X. Laskibar, J.M. Lekuona & |. Salcedo, 6374 IS,
BlO-Fungi 4694. Aya, Altxerri stream, 180 m, 30TWN6888, on
wood very wet, 21-XI-1992, J.L. Albizu, J. Huarte, X. Laskibar, J.M.
Lekuona & |. Salcedo, 6420IS, BlO-Fungi 4695.
A species which is mainly found in tropical areas; until now it was only
to be found in Europe in greenhouses and mines (cf. Ginns, 1980:1581;
Ryvarden & Gilbertson, 1993:253). The specimens studied grow in open
places (out of doors), which are always very humid. We will add that, in
1992, the species has also been found in Bayonne (France) by David &
al. (pers. com.).
The sulphur-yellow colour of the basidiocarp and the hymenophore
poroid with 9-10 pores per mm makes the identification of the species
in the field easy.
The species is also usually easy to recognize because of the presence of
numerous cilindric, long cystidia, thickly walled and strongly
encrusted in the upper part, embedded in the trama or projecting
slightly above it (fig. 3K); also by the ellipsoid basidiospores, about
2.5-3 x 1.6-2 ym, since both are diagnostic characters of this taxon.
Fig. 3. A-Acanthobasidium phragmitis, BlO-Fungi 4693, pleuroacantho
basidiole. B-ldem, gloeocystidia. C-ldem, pleuroacanthobasidioles and
spores. D-ldem, shriveled and deformed spores. E-Auriporia aurulenta,
BlO-Fungi4690, hymenium with cystidia. F-ldem, hymenium with
cystidia. G,H-Sarcodontia crocea, BlIO-Fungi 4691, spores and basidium.
|,J-Auriporia aurulenta, BlO-Fungi 4690, spores. K-Flaviporus
brownei, BlO-Fungi 4695, hymenium with cystidia. L-Sarcodontia
crocea, BlO-Fungi 4691, hymenium.
6
Sarcodontia crocea (Schwein.:Fr.) Kotl. Ceska mykol. 7:117
ClYss)
= S. setosa (Pers.) Donk
GUIPUZCOA: Hernialde, 250 m, 30TWN7478, on Malus sp., 25-X-
1992, J. Huarte, 62691S, BlO-Fungi 4691.
As indicated by Breitenbach & Kranzlin (1986:168) and Jilich
(1984:171) this species is of a wide distribution but with a very
concrete habitat as it usually grows on apple trees, especially old ones.
The resupinate basidiocarp, or stratified on vertical substata,
hymenophore hydnoid with 5-7 mm long aculei, are the characteristics
to be emphasized, as well as the sulphur-yellow colour and intensely
nasty smell. When dried it loses some of its colour and smell .
The specimens studied agree with the description given by Eriksson &
al. (1981:1275); nevertheless we underline the fact that its spores are
subgloboses, smooth, with somewhat thickened walls, and usually with
an oil-drop, and about 4.8-5 x 4 um (figs. 3G,3H,3L).
ACKNOWLEDGEMENTS
| would like to thank all those who have helped me compile the
specimens. And A.J.LOpez-Quintana and M.T.Telleria for their help in
checking the manuscript.
REFERENCES
BOIDIN, J.,_ANQUETIN,P., CANDOUSSAU,F., GILLES,G. & HUGUENEY,R.
(1985). Contribution a la connaissance des Aleurodiscoideae a spores
amyloides (Basidiomycotina, Corticiaceae). Bull. Soc. Mycol. France
101. (4)(333-360
BREITENBACH,J. & KRANZLIN,F. (1986). Fungi of Switzerland. Non
gilled fungi (Heterobasidiomycetes, Aphyllophorales, Gastromcetes.
vol. 2 Verlag Mykologia. Lucerne.
DAVID,A., TORTIC,M. & JELIC,M. (1974). Etudes comparatives de deux
espéces d'Auriporia. Bull. Soc. Mycol. France 90 (4):359-370.
DONK,M.A. (1960). The generic names proposed for Polyporaceae.
Persoonia 1 :173-302.
ERIKSSON,J., HJORTSTAM,K. & RYVARDEN,L. (1981). The Corticiaceae
of North Europe. vol. 6. Fungiflora. Oslo.
GINNS,J. (1980). The genus Flaviporus Murrill (Polyporaceae) Can. J.
Bot. 58 :1578-1590.
JULICH,W. (1984). Die Nichtblatlerpilze, Gallertpilze und Bauchpilze.
Gustav Fischer Verlag. Sttugart. 626pp.
7
RYVARDEN,L. & GILBERTSON,R.L. (1993). European polypores. part 1.
Fungiflora. Oslo.
SALCEDO,|. (1989). Catélogo comentado de los Aphyllophorales
(Basidiomycotina) del territorio historico de Alava. Tesis doctoral.
Universidad del Pais Vasco/EHU.
SALCEDO,|. & TELLERIA,M.T. (1992). Contribucion al conocimiento de
los Aphyllophorales del Pais Vasco. |. Bol. Soc. Micol. Madrid 16 :53-
60.
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MY COTAXON
Volume L, pp. 9-18 January-March 1994
CONTRIBUTION TO THE LICHEN FLORA OF TRANSKEI
C.M. THOMAS
Department of Education, Umtata, Transkei. South Africa
R.B. BHAT
Department of Botany,University of Transkei ,Private
Bag XI, Umtata, Transkei, South Africa
ABSTRACT
Fourteen species of lichens which have been discovered
as new to Transkei, are described. The result of the recent
investigations are based on the collections made from nine
districts of this region. The physical selective forces limit
the period of acquisition of resources needed for the growth.
Frequent changes in the elements of climate, such as
unexpected cold wave, irrespective of the season are
contributing factors to the relatively small number of
species. Lichen flora of Transkei have been unexplored for
a long time. Interesting and new species are expected from
this region. No lichen community is reported as endangered.
Changes in the lichen flora cannot be assessed, as there is
no previous comprehensive record of lichens in Transkei.
INTRODUCTION
Lichenological exploration of the Transkei region
started in 1953. Collections made by Almborn from Port St.
Johns and Flagstaff were the first, to reveal the existence
of interesting endemic taxa and ubiquitous species in this
area (Almborn 1966). Distinct foliicolous Lichen specimens
from Dwesa Nature Reserve are illustrated by Brusse (1992).
The present account is a contribution to the lichenological
exploration of Transkei. In all, 500 collections were made
belonging to 115 species. Some of them have been identified
and others are under the process of identification. The
identified specimens are new records for the Transkei region
and described in this paper. In spite of all previous
investigations many and interesting parts of Transkei remain
unexplored as far as the lichen flora is concerned. Among
the species recorded as new for Transkei come under fruticose
5 and foliose 9.
10
MATERIALS AND METHODS
The following account is based on the collections made
by the authors in 1992 from nine districts of Transkei, South
Africa. The materials are placed in the authors herbarium and
the representative set deposited in the herbarium of the
University of Transkei (KEI) for future reference. The
following number codes are used to indicate the collecting
localities and their altitudes.
Port St Johns Psd 1 5= 2.9 Pl
Hululeka Nature Reserve HNR 2 200 M
Libode LBD 3 930 M
Butterworth BTH 4 680 M
Encgobo EGO 5 1200 M
Langeni LGI 6 900 M
TsolorTLony 1300 M
Mt. Frere MFR 8 1200 M
Umtata UTA 9 800 M
The specimens were collected from different localities
(see map of Transkei), dried and preserved by customary
method. Visits to the localities were made at different
seasons. Morphology of the upper and the lower surfaces of
the thallus, rhizine occurrence and the fruiting bodies were
examined for identifications. Macrophotographs were taken
with a cannon camera using close up view lens X 6.
ECOLOGY
Growth of most lichens is controlled by specific
requirements like moisture, light and temperature. These
factors together with altitude effectively restrict their
distribution. The physical selective forces limit the period
of acquisition of resources needed for growth.
Transkei is between 27°-30° E longitude and 30°-33° §
latitude, has an area of 58024 Ha. The ecologic - geographic
variations are from coastal region hilly midland to
mountainous interior with sub tropical climate with hot wet
summer and warm dry winter. The interior is similar to coast
but cooler with increasing height. The average daily minimum
temperature of the coldest month (July) is 7° c and average
daily maximum temperature of hottest month (January) is 28°
c. Average rainfall in summer season is 600 mm. although it
varies greatly from place to place. Average rain fall in
winter is 200 mm. Towards the end of the winter season, this
peninsular region experiences strong gale blowing from south
polar region. Relative humidity is approximately 40% in the
interior and it varies from 60% = 70% > at’ the’ coast “and
midland. The altitude varies from sea level to 2000 nm.
Habitat ranges from dry mountainous area, grass topped bare
hills, humid forests and wild coast supporting varied lichen
fiora;
Map of Transkei
12
Description of the species are based exclusively on materials
studied by the authors. Many taxonomically or
phytogeographically poorly investigated species, are not
included in the list for the time being. The following
collector’s abbreviation is used (R.B.B - R.B. Bhat) for the
specimens.
R.B.B. 101. Parmelia(Rimelia) reticulata Tyl.
Corticolous, foliose, upper surface of the thallus is green,
lower surface brown along the tip and black towards the other
regions. Apothecia not very common. Densely rhizinate on its
lower surface. Width of the thallus 8-10 mm. It is commonly
found in Umtata.
R.B. B. LO2. skancnoriavparvetina ibs john ay Pin
Corticolous. This is a bright orange red foliose species.
Thallus turns green when it is wet. Sparsely rhizinate on
appressed parts of the thallus. Round apothecia is sessile
and adnate. Apothecial discs are orange red in colour. Grows
on the fresh bark of the shady trees along Umtata river bank
behind St.Bed’s College. It prefers shady places with high
humidity at 400 m - 600 m MSL. Width of the thallus 3-4 mn.
It is a rare species.
R.B.B 103. Parmelia(Flavoparmelia) soredians Nyl.
Corticolous, foliose, upper surface of the thallus yellowish
green, lower surface black, moderately rhizinate on appressed
parts of the thallus. It is commonly found on the fresh bark
of maple trees near stadium of the Transkei university campus
Umtata. Width of the thallus 2-5 mm. Apothecia not common.
R.B.B. 104. Parmelia(Punctelia)borreri (Sm.) Krog.
Corticolous, foliose, upper surface of the thallus is green
lower surface is brown and black. Short rhizines occur.
Found on dry/fresh bark of shady trees along the bank of
Umtata river on the lower side of St. Bedes College. Width of
the) thallus: (5=72 aim.
R.B.B. 105. Dirinaria picata (Sw.) Aem. & Shear
Corticolous, foliose, upper surface of the thallus is grey,
lower surface black. Thallus firmly attached to the
substratum. Grows on dry bark of the trees along the bank of
Umtata river. 5-6 cm in diameter. Width of thallus 2-4 mn.
Plate 1- General aspects of Parmelia and Heterodermia spps.,
showing different forms of lobes and apices.
a- Parmelia (Punctilia) borreri b- Heterodermia diademata
c- Parmelia(Rimelia)reticulata d- Parmelia(Xanthoparmelia)
mutabilis e- Parmelia(Flavoparmelia)soredians f- Parmelia
(Parmotrema) austrosinensis Magnification x 1
is
14
R.B.B. 106. Heterodermia diademata (Tayl.) Awasthi
Corticolous, foliose, upper surface is greyish white, lower
surface white, and rhizinate. Apothecia is rare. Width of the
thallus 1-2 mm. This genus is known before only from tropical
America, Africa and Philippines and New Gunea.
R.B.B. 107. Parmelia(Parmotrema) austrosinensis Zahlbr.
Corticolous, foliose, upper surface is green towards the
Older parts of the thallus and grey towards younger
parts.Lower surface rhizinate, brown. Occur on the fresh
bark of citrus trees in the orchards of Agriculture college,
Tsolo. Grows to 3-5 cm in diameter. Width of the thallus
varies from 5-12 mn.
R.B.B. 108. Parmelia(Xanthoparmelia)mutabilis Tayl.
Saxicolous, foliose, upper surface olive green, lower surface
brown, rhizinate. It is highly evolved group in southern
Africa (Hale Jr.1972). Average width of the thallus 3-5 mn.
R.B.B. 110. Physcia stellaris (L.) Nyl.
Corticolous, foliose, upper surface grey, glossy, grows on
fresh bark of shrubs along Umtata river behind St. Bedes
College.
R.B.B. 111. Teloschistis exelis (Michx.) Vain.
Corticolous, fruticose. Grows on the fresh bark of jacaranda
trees, garden rose plants at Buntigville, Umtata General
hospital. Prefers shrubs. Considerable variation in thallus
colour is observed from yellow to orange red to grey or
greyish white. Highly branched thalli form tufts of about 5
cm diameter. Long cylindrical very slender erect lobes.
Thallus lobes are 3-4 cm long and 0.5-1 mm wide,narrows
towards the tip. Tips of the thallus end like a spine.
Dichotomously branched. Apothecia frequent. Apothecial discs
are orange red in colour and flat. They are lateral or
terminal. No hairs at the margin of the apothecia. Orange red
fruiting bodies are held on stalks.
R.B.B. 113 Teloschistis hypoglaucus (Nylk.) Zahlbr.
Corticolous, fruticose lichen with foliose thallus as seen in
lichens of the Himalayas (Poelt & Obermayer 1991). Thallus is
divided in to many lobes. Grows to 1-2 cm in diameter.
Thallus is ash grey or grey in colour. Lower surface is
reticulately veined. Apothecia are numerous and they are
lateral or terminal. Apothecial discs are orange red as in
Plate 2- Upper surface of thalli of fruticose and foliose
lichens. a- Teloschistes hypoglaucus with apothecia (arrow)
b- Xanthoria parietina c- Dirinaria picatata
d- Usnea undulata e- Ramalina aspera with apothecia (arrow)
f- Ramalina celastri thallus with apothecia (arrow)
15
16
Xanthoria and concave, 1-2 mm in diameter. Hairy structures
along the margin of the apothecia and on the thallus.
Apothecia are held on short stalks. Grows on fresh bark of
shady trees along gravel road to Zingiza. Occurs only in
southern hemisphere, South America, South Africa and Australia
(Almborn 1989).
R.B.B. 115) Usnea undulata (Stirt.)
Corticolous, fruticose, green, erect, or pendulous
dichotomously branched thallus. Radially symmetrical. The
thallus is attached to the substratum at one point. Grows on
fresh bark on trees as well as on dry branches. Varying in
height from 3-8 cm. Occurs very commonly in Hululeka Nature
reserve, Engcobo, and Libode.
R.B.B. 118. Ramalina celastri (Spreng.) Krog. & Swinsc.
Corticolous, fruticose, thallus is short with broad undivided
base. Thallus flat and ridged. Olive green in colour.
Bilaterally symmetrical,firmly attached to the fresh bark of
trees. The base of the thallus penetrate deep in to the
cortex of the host plant. Thallus is branched. Apothecia are
abundant and adnate, apical or sub apical or laminal varying
in height from 1-8 cm. Width of the thallus varies from 3-10
mm. In some cases the lamina of the thallus has foliatious
extensions. R.celastri is one of the most common species in
Transkei. It is found to be widely distributed on branches
which are exposed to light,mostly upper part of the trees and
usually not observed to towards the base of the host plant.
Photophilous.
R.B.B. 120. Ramalina aspera Ras.
Corticolous, fruticose broad and flat thallus,bilaterally
symmetrical, branched, short, 1-3 cm long, 3-5 mm broad. The
basal part is un-branched. Apothecia sub apical or sometimes
laminal or marginal or at random held on very short stalk.
Apothecial disc flat. Occurs more frequently on exposed
branches of trees,therefore tolerant to high intensity of
light. It is also observed on artificial habitats like parks
and town avenues on fence posts roof tops. They occur more
frequently on well exposed branches compared to lower
branches which are shaded. Photophilous.
CONCLUSION
Fourteen lichens new to Transkei are investigated and
recorded. They are mostly common or wide spread species,
which indicate that most part of the ‘Transkei and
scientifically important areas are unknown from. the
lichenological point of view. The coastal region represents
a community dominated by different species of Ramalina
together with foliose and crustose lichens. As the
population of Ramalina increases on the host plant the
vegetative growth of that particular part decreases and
Slowly dies off giving a clue that it may be semi-parasite.
T7
Further this genus shows heliotropic tendency. The altitude,
strong gale from polar region, frequent changes in the
elements of climate are contributing factors to the
relatively small number of species in this region. Some
areas are so inactive that vegetation does not have the
opportunity to become established. In areas where vegetation
occurs it can appear as strips and patches. Some types of
lichens especially foliose and fruticose establish themselves
by becoming intertwined with host plant material. Some
fruticose lichens which have foliose thallus, is a special
feature noted in Transkei. Similar observations have been
made in some lichens in the Himalayas (Poelt & Obermayer,
1991). Lichens can be used as biological indicators because
of their sensitivity to environmental pollution. Scientific
research regarding the role of lichens as monitors of
environmental pollution began from the middle of the century.
As there is no previous comprehensive records of lichens from
this regions, the rate of pollution and changes in lichen
communities could not be assessed and established. No
particular lichen community is reported as endangered by any
agency. Any type of environmental change may have an impact
on the lichen population. Growth in human population and
industries in the city of Umtata and other towns may increase
pollution and influence the growth of lichens and may result
in the disappearance of some lichen species. Further future
investigations on the lichenological studies may throw more
light in this direction.
ACKNOWLEDGEMENTS
We are grateful to the Department of Forestry, Transkei,
South Africa, for the necessary permit to collect the living
materials from the forests and nature reserves. Our thanks
are also due to Dr. Darrell J. Weber for the ' valuable
suggestions and corrections to the manuscript.
REFERENCES
Almborn, O. 1966. Revision of some lichen genera in
southern Africa I. Botaniska Notisier 119
OF ase Sy. sa7.0 slo,
Almborn, O. 1989.Revision of the lichen genus Teloschistes
in central and southern Africa. Nord. J. Bot.
8(5): 521-537.
Brusse,F.Ai, Dickinson, -C.H..1991.)Asnew foliicolous
species in the lichen genus Porina (Porinaceae,
Pyrinulales) from southern Africa. Mycotaxon 42:
Sidi sO Os
Brusse, F. 1992. Gomphillaceae (Lichens). A new species of
Bullatina from the Transkei wild coast. Bothalia
22 (1): 44-46.
18
Elix, J.A. & Johnston, J. 1988. New species and new reports
of Flavoparmelia (lichenized Ascomycotina) from
southern hemisphere. Mycotaxon 33: 391-400.
Hale Mason, E. Jr. 1972. New species of Parmelia section
cyclocheila in southern Africa. Bryologist 75
(3): 342-348.
Poelt, J. & Obermayer,W. 1991.Beitrage zur Kenntnis der
Flechten Flora des Himalaya II .Die Gattung
Byonora (Lichenes, Lecanoraceae) Zugleich eine
revision aller Arten. Nova Hedwigia 53(1): 1-26.
MY COTAXON
Volume L, pp. 19-25 January-March 1994
BATTARREOIDES DIGUETII (GASTEROMYCETES, TULOSTOMATALES) IN
SOUTHERN AFRICA
J.C. COETZEE and A. EICKER
Department of Biological Sciences, Peninsula Technikon,
P.O. Box 1906, Bellville, 7530, Republic of South Africa
Department of Botany, University of Pretoria, 0001,
Republic of South Africa.
SUMMARY
Two recent collections of Battarreoides diguetii from
Botswana and South Africa confirm the occurrence of this
fungus in southern Africa. The southern African material
is described and its spores are compared with those of
Batarrea stevenii.
KEY WORDS: Battarreoides diguetii; Batarrea stevenii;
southern Africa; scanning electron microscopy.
INTRODUCTION
When Bottomley (1948) recorded Battarreoides diquetii
(Pat. & Har.) R. Heim & T. Herrera [as Batarrea digueti
Pat. & Har.] from the northern Transvaal in South Africa,
she had little doubt that her diagnosis was correct, even
though this species was, until then, known from, and
believed to be confined to North America only (Cunninghan,
1944). Scepticism regarding the occurrence of this fungus
in Africa persisted, however, and in his authoritative
Gasteromycete overview, Dring (1973) for example, still
cited Battarreoides as an endemic North American genus, a
view also upheld in Hawksworth et al. (1983). The disjunct
distribution pattern created by the African material,
which comprised of a single specimen only, and the
uncharacteristic photograph in Bottomley (1948) might have
contributed to this scepticism. The recent collection of
three Battarreoides specimens from the Kalahari region in
Botswana and South Africa was of particular significance,
therefore, since it finally confirmed the southern African
distribution of this genus. The Kalahari material, as well
as the northern Transvaal specimen (Bottomley, 1948),
largely agree to earlier descriptions of Battarreoides
Giguetii (Patouillard & Hariot, 1896; Rea, 1942; Herrera,
1953) and these authors believe that the southern African
20
material should be assigned to that species.
MATERIALS AND METHODS
The description that follows is based on dried
herbarium material kept in the South African National
Collection of Fungi (PREM). Only three southern African
collections, comprising four specimens, exist. Kreisel
(1967) and \Miller)))& Miller: ((1988))\.were (followed) > in
determining the blueing reaction of the glebal hyphae in
aniline blue/lactophenol. For electron microscopy, glebal
tissue of B. diguetii was affixed onto copper stubs,
coated with gold ina Polaron sputter coater and examined
with a Jeol 840 scanning electron microscope. For
comparison, spores of two Batarrea stevenii collections
(PREM 28261 and PREM 49215) were examined in the same way.
Spore sizes were measured directly from the scanning
electron micrographs, but capillitium measurements were
obtained from material mounted in colourless lactophenol
and observed through a| Reichert-Jung Polyvar microscope.
Colour names and codes appearing in brackets’ refer to the
closest colour equivalents in the "Methuen Handbook of
Colour" (Kornerup & Wanscher, 1978).
DESCRIPTION
Battarreoides diguetii (Pat. et Har.) R. Heim et T.
Herrera,! Anwiinst «biol «Mex. wo2 em SU (LOGI.
= Batarrea Digueti Patouillard et Hariot,
Journ. /de Botan due!) 0015 est 2 5 pon ek
(1896).
= Battarraeastrum Digueti (Pat. et Har.) R.
Heim et T. Herrera, Rev. de Mycol. 25(3-4):
2 Die bn L9 GO.)
= Battarrea Griffithsii Underwood, Bull. Torrey
Bot. Olub: 28-4440) opty o3 7k OOi ie
= Battarrea Digueti forma minor Lloyd, Myc. Writ.
TAS) ah S aD Lees ty noe 3 SG Oo aL
= Battarreoides potosinus T. Herrera, An. Inst.
Biol Mex vu243 (47 =A26,,(01953)).
Basidiocarp stipitate. Peridium attached more or less
centrally on the stipe apex, campanulate, 30-51 mm wide,
up to, 30), mm high, Vbut only;up. to 12. smnmiseparating che
peridial apex from the point of attachment to the stipe. A
narrow, protruding, membraneous margin demarcates’ the
junction between the convex upper and concave bottom
peridial halves (zone of circumscissile dehiscence in
Batarrea). Exoperidium creamish white (yellowish white;
442), thin, brittle fugacious,,-virtually absent onwsome
(usually weathered) specimens. Endoperidium concolourous
with exoperidium, smooth, texture tough and springy,
dehiscing by means of pores mostly arranged ina circle
along the peridial circumference, a few millimetres above
the junction of the upper and lower peridial halves. Stipe
zt
yellowish cream (pale yellow; 4A3) or blackish brown
(although more brownish towards the base), up to 27,5 cm
long, 9-12 mm wide, cylindrical but attenuating towards
the base, hollow, smooth to squamose, may be deeply
sulcate, lacerating into large linear, fibrous scales,
more so towards the base, sometimes contorted, the
sulcations then spiralling along the length of the stipe.
Gleba cobwebby, dark brown (dark brown; 7F8), adherent to
the base of the endoperidium, comprised of elaters, spores
and hyaline to very faintly pigmented hyphae (both thin
walled and skeletal), 2-7 um in diameter. Thin-walled
hyphae abundant, comprising single hyphae to amorphous,
tissue-like mycelial aggregates, sparsely septate,
occasionally branched. Skeletal hyphae (capillitium) thick
Wotltece (Up icOmlls/> Lum rarely branched.) “apparently
aseptate, cyanophilic, arising from thin walled hyphae.
Elaters 2,5-8,75 wm in diameter (mostly varying between
4,5-6,25 wm), consisting of very short to longer fragments
usually not exceeding much more than 160 yum in length
(although lengths of up to 210 um have been measured),
occasionally branched, often with tapering, rounded ends.
Elater walls hyaline and thin but with numerous annular
(occasionally spiral), wall thickenings, yellowish brown
in colourless lactophenol and positively cyanophilic upon
heating in aniline blue/lactophenol. Basidia: immature
stages not preserved to record these cells. Basidiospores
dark rusty brown (dark. brown; 7F8).,,. globose’ to’ broadly
Nata e-Oy oO. LM LOnGsand 4)6—-576 Uso rOag.! Very sSnoncly
pedicelled (0,5 um), appearing pitted under the light
microscope’ but with a aqistinctiy reticulate wall
sculpture on the electron micrographs. Volva not observed.
Habitat: Arid soil; sand dune; termite mound.
Distribution in Africa: northern Transvaal and north
western Cape, South Africa; central Botswana.
Distribution elsewhere: North America.
Teones: Patouillard & | Hariot))(1896),, White (1901), \Lioyd
(29063. PLOVaS NT CLO23 ie REAM GLO d 2) BOCCOmMLey i \(LO4Syr,
Herrera (1953), Heim & Herrera (1960).
Material examined: SOUTH AFRICA: Botanic Reserve near
Messina, northern Transvaal, on termite mound, I.B. Pole
EBVans. |S PREM. 20459, August; | 19257. Groblershoop, north
western Cape Province, on sand dune, M.C. Moolman, PREM
494325) March 1989. BOTSWANA: Approximately 8 km West of
Tsutswa pan (approx. 63 km West of Hukuntzi), Kgalagadi
SIisteict, On sanay soll, 1G. D. .Oubpert (PREM) 51435 7)'13 .97.89.
DISCUSSION
Macroscopically, the monotypic genus Battarreoides
differs from Batarrea by the endoperidium which dehisces
by multiple pores instead of circumscissilely. The most
Significant microscopic difference between the two genera
lies in the relative lengths of the elaters, which in
Batarrea seldomly exceed 80 um, while lengths of between
100-200 wm are common in Battarreoides (Rea, 1942; Miller
& Miller, 1988).
22
Although the photograph of Battarreoides diguetii in
Bottomley (1948) depicts a specimen without peridial
perforations, re-examination of Bottomley’s material (PREM
20459) did reveal distinctive perforations in that part of
the peridium not visible in the photograph. Examination
and careful comparison of ‘all the southern African
specimens convinced uS'/5Of Sthexr conspecificity and
position in the genus Battarreoides.
The southern African Battarreoides specimens do not
differ Significantly from earlier descriptions and
illustrations of B. diguetii (Patouillard & Hariot, 1896;
White, 1901; Lloyd, 1906; Lloyd, 1923; Rea, 1942; Herrera,
1953; Heim & Herrera, 1960) and we have little doubt that
they represent the same species. The peridial perforations
in our specimens are largely restricted to a circular
arrangement and, with one exception, are not scattered
over the peridial surface as reported by Patouillard &
Hariot (1896) and Herrera (1953). This seems to bea
variable characterisric, however, as a Circurar
arrangement had also _been reported for a number of
specimens examined by Herrera (1953)..5 MOsStasrautcnons
describe the gleba as pulverulent and not cobwebby as
interpreted by Lloyd (1923) and ourselves. This character
also seems to vary, and is speculated to be a function of
the’ age* ofthe fruitang body at the “time of “collection,
The only major difference between the southern African and
North American material appears to be the absence of a
volva,in the former. Thais, however,.1s a’ structure tac
could easily be overlooked by the inexperienced collector
and from earlier illustrations it is evident that the
volva is not always included in herbarium specimens
(Lloyd, 1923; Rea, 1942; Heim & Herrera, 1960). All the
southern African collections were made by non-mycologists
and the absence of the volva is therefore most likely to
have been caused by collector ignorance.
The spore ornamentation of B. digueti has been
interpreted as warted, reticulated or perforate (Rea,
1942), obscurely verrucose (Patoulllard & :Hariot, 1896)
and finely bristly (Herrera, 1953). Under the SEM, spores
of the southern African collections were, however,
distinctly” retictilatere(hrgures ila sande ab)7 By this
characteristic alone, the Battarreoides specimens were
clearly distinguishable from Batarrea stevenii which had
spores of a more verrucose nature (Figures 1c and 1d). The
reticulate spore ornamentation, lack of circumscissile
dehiscence and long elaters thus form a combination of
characteristics which distinguish Battarreoides from
Batarrea. This”1s* in tcontlict with) the results ors sagoom
study by Liu and Liu (1983), however, who reported and
Fig. 1. a-b. Basidiospores of Battarreoides diguetii.
c-d. Basidiospores and elaters of Batarrea
stevenii.
24
illustrated reticulate spores, closely resembling those of
our Battarreoides diguetii specimens, for Batarrea
stevenii. This apparent discrepancy requires an
examination of the sporophores used in their study for
elater length and circumscissile dehiscence. Tt ce
probable that the spores described by Liu and Liu (1983)
were not those of Batarrea stevenii.
ACKNOWLEDGEMENTS
The co-operation of Alice P. Baxter, curator of PREM,
and’ the assistance of Prof. J. Coetzee, Mr. C. van der
Merwe and Mr. A. Botha of the electron microscopy unit,
University of Pretoria, is gratefully acknowledged.
Professor Orson Ki. Miller of Virginia Polytechnic
Institute and State University is thanked for critically
reviewing an earlier draft of this manuscript. This study
was partially funded by a FRD grant to the junior author.
REFERENCES
Bottomley, A.M. (1948). Gasteromycetes of South Africa.
Bothalia 4(3): 73-810.
Cunningham, G.H. (1944). The Gasteromycetes of Australia
and New Zealand. Dunedin: John McIndoe.
Dring, D.M. (1973). Gasteromycetes. pp. 451-478. In
Ainsworth et al., (eds): The Fungi: An advanced
treatise. Vol 4B. New York: Academic Press.
Hawksworth)«(D. Li /{Sutton Bwie. To \Aansworth, G.Cly (L983),
Ainsworth and Bisby’s Dictionary of the Fungi, 7th
ed. Kew : Commonwealth Mycological Institute.
Herrera, T. (1953). Un hongo nuevo procedente del Estado
de San Luis Potosi Battarreoides potosinus. An.
Inst. Biol. Mex. 24: 41-46.
Heim, R. & Herrera, T. (1960). Nouvelles contributions a
la flore mycologique mexicaine. Revue de Mycologie
ZO mi clo) 236
Heim, R. & Herrera, T. (1961). Una nueva especie de
Podocrea y una nueva combinacion el nombre de un
hongo Mexicano del genero Battarreoides. An. Inst.
Biol.) Mex. 322)°29-3:1;
Kornerup, A. & Wanscher, J.H. (1978). Methuen handbook of
colour. London: Methuen.
Kreisel, H. (1988). Taxonomisch-Pflanzengeographische
Monographie der Gattung Bovista. Beihefte zur Nova
Hedwigia 25: 1-244.
Liu, Be) &) baw Yi -HS 4983), Scanning, electron :;microscopy
of basidiospores of Battarraea stevenii. Mycologia
VD e OL = Ob.
Lloyd, C.G. (1906). Tylostomeae. Mycological Writings 2:
Se Zor.
Lloyd, .CaG.., (1923)). ‘Battarrea Digqueti,.from) Ivan iM.
Johnston, Collected:San Nicholas Bay, California.
Mycological Writings’ )7'(3 }swLl74—1175).
Miller, O.K. & Miller, H.H. (1988). Gasteromycetes.
25
Morphological and development features with keys to
the orders, families and genera. Eureka: Mad River
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Patouillard, M.M. & Hariot, P. (1896). Liste des
champignons recolte en Basse-Californie par M.
Diguet. Journal de Botanique 10: 250-251.
Rea, P.M. (1942). Fungi of southern California. 1.
Mycologia 34: 563-574.
White, Vis. (1901). The Tylostomaceae of North America.
Bulletin of the Torrey Botanical Club 28: 421-444.
MY COTAXON
Volume L, pp. 27-46 January-March 1994
TYPE STUDIES IN THE POLYPORACEAE 25
SPECIES DESCRIBED FROM JAPAN BY R. IMAZEKI AND A. YASUDA
BY
TSUTOMU HATTORI
Forestry and Forest Products Research Institute, PO. Box 16, Tsukuba Norin,
Kenkyu, Danchi-Nai, Ibaraki, 305 Japan
and
LEIF RYVARDEN
Department of Botany, University of Oslo, P.O.
Box 1045, Blindern, N-0316 Oslo, Norway
ABSTRACT
The types of 32 polypores described by R. Imazeki and A. Yasuda have been
examined. 17 species are accepted while 15 are placed in synonymy with already
described species. The following new combinations are proposed: Melanoporia
castanea (Imaz.) Hatt.& Ryv., Antrodiella gypsea (Yas.) Hatt. & Ryv., Perenniporia
japonica (Yas.) Hatt. & Ryv., Perenniporia minutissima (Yas.) Hatt. & Ryv. and
Trichaptum parvulus (Yasuda) Hatt. & Ryv.
INTRODUCTION
The polypore flora of Japan is incompletely known and the latest comprehensive list
is that of Ito (1955). However, numerous resupinate species were omitted while
many species have later been reported from the country, see for example Aoshima &
Kobayashi (1983) and Furukawa, Abe & Neda (1983). Further, the nomenclature in
Ito's list is outdated, making it difficult to use the list.
The flora of North American polypores (Gilbertson & Ryvarden 1986 & 1987) gave
an updated checklist for the North American continent using the same nomenclature
as in Europe (see Ryvarden & Gilbertson 1993). These publications have made
comparative studies much easier since the names used in the two areas now are in
consensus,
From Ito's list and more popular floras like those of Imazeki & Hongo (1965, 1989),
it is evident that Japan has many polypores in common with both Europe and North
America. Some of them have been named anew in Japan while others are cited with
synonyms. Thus, we felt it necessary to make up a critical checklist for the polypores
of Japan as a first preliminary step towards a complete flora. During the compilation
of names of polypores recorded from Japan, it became clear that a number of types
28
ND000D 0
10 wm
Nie
Fig. 1. Melanoporia castanea A) section of basidiocarp, B) basidiospores, C)
basidia, D) contextual generative hyphae, E) contextual skeletal hyphae. A,D and E
from the type, B and C from TFM-F-15770.
Ne)
had to be examined to ascertain the status of a number of names which more or less
had slept in oblivion since their publication. The results of these studies are
published below to avoid excessive taxonomic discussions and descriptions in the
forthcoming checklist.
The two authors cited in the title are the only Japanese mycologists who have
published new polypores from the main land of Japan. Bonin Island has a very distinct
mycoflora being situated 1,000 km SE of Japanese main islands, and its polypores
will be treated in a series of separate papers. Thus, we felt it natural to treat R.
Imazeki and A. Yasuda species in one paper.
C. G. Lloyd described many species from Japan based on collections from Yasuda.
The results of studies on the types of these species have already been published in
previous parts of this series. The reader is referred to Ryvarden (1991) for a
complete list of previous type studies and a list of all species treated in the series up
to 1991.
In the following, species of both authors are listed alphabetically according to specific
epithet. For each species the place of publication is given with the type locality and
the herbarium in which the type is deposited. The herbaria are indicated with the
abbreviations as given in Index Herbariorum, 7th edition.
K. Aoshima proposed many names for new polypores from Japan. However,
unfortunately he overlooked the rules of the International Code of Botanical
Nomenclature. Thus, all his new names, mostly as nomina nuda, must be rejected as
invalidly published for different reasons.
If the species is accepted, it is cited in its proper genus with a reference to a modern
description. If none exist, the species is described here. The basionyms and the
accepted names are printed in bold face.
LIST OF SPECIES
albicans, Grifola Imaz., J. Jap. Bot. 19:386, 1943.
Yabukawa, Iwate Pref., Japan, 29 Sep. 1938, leg. K. Ito (lectotype selected here,
TNS-206974).= Grifola frondosa (Fr.) S.F.Gray The type specimen represents only a
pale form of the widespread G. frondosa. Imazeki (1943) concluded that it is distinct
with its very wide context hyphae, but such wide hyphae are also seen in many
specimens of G. frondosa.
calvatioides, Polyporus Imaz. J. Jap. Bot. 16:269, 1940.
Summer, 1917, leg. J; Umemura (Type of Calvatia versispora Lloyd, BPI).
= Laetiporus versisporus (Lloyd) Imaz.
P. calvatioides was proposed as nomen novum (for use in the genus Polyporus) by
Imazeki because the combination Polyporus versisporus Lloyd was an illegitimate
homonym of Polyporus versisporus Pers. 1825.
castanea, Fomitopsis Imaz., Bull. Gov. Forest Exp. St. Tokyo 42:1, 1949.
Kitasaku, Nagano, Japan, 9 Oct. 1947, leg. R. Imazeki (type, TFM - F-346; isotype in
TNS).
=Melanoporia castanea (Imaz.) Hattori & Ryvarden comb. nov., basionym as
30
Fig. 2. A-C: Daedaleopsis conchiformis A) basidiocarps, B) section of basidiocarp,
C) pores. From the type. D-I: Daedalea dickensii D) pores, C) basidiospores, F)
basidia, G) contextual generative hyphae, H) contextual skeletal hyphae, 1)
contextual binding hyphae. D, G-I from the lectotype; E and F from TFM-F-16200.
31
cited above.
Basidiocarp sessile, occasionally resupinate, perennial; pileus triquetrous, ungulate
or irregular, up to 30 cm long, up to 15 cm thick, margin thick and dull, pileus surface
chestnut brown and velutinous in young specimens, almost black and glabrous in old
specimens; tubes multizonate, purplish brown, pore surface dark umber to purplish
brown, pores circular, 5-6 /mm; context fibrous-corky, purplish brown.
Hyphal system dimitic, contextual generative hyphae hyaline, 1.5-4 um wide, with
clamp connections; contextual skeletal hyphae olivaceous brown in KOH, 3-5.5 um
wide, thick to almost solid, unbranched to occasionally branched; tramal generative
hyphae hyaline to pale yellow, 1.5-3 um wide, with clamp connections; tramal
skeletal hyphae olivaceous brown in KOH, almost straight, thick to almost solid, 3-5
uum wide; cystidia none. Type sterile, so following descriptions of basidia and
basidiospores were made based on TFM-F-15770. Basidia clavate, 15-20 x 4.5-5 um,
4-sterigmate, with a basal clamp, basidiospores long ellipsoid, hyaline, IKI-, 4-5 x
1.8-2.5 um (Fig. 1).
Remarks: This species is closely related to Melanoporia nigra (Berk.)
Murr.(basionym Polyporus nigra Berk., K, BPI!) with its purplish brown context and
brown rot. Thus, Imazeki & Hongo (1989) reduced it to a synonym of M. nigra.
However, M. nigra has a consistently resupinate basidiocarp while M. castanea
usually has a well developed pileus. Besides, the context hyphae of M. nigra are
much wider than those of M. castanea.
cercidiphyllum, Cryptoderma Imaz., Bull. Gov. Forest Exp. St. Tokyo 42:2,1949.
Shibetsu, Hokkaido, Japan, 23 July 1941, leg. R. Imazeki (type, TNS-212095).
= Phellinus conchatus (Pers.:Fr.) Pat. as already pointed out by Aoshima (1983).
citrinum, Cryptoderma Imaz., Bull. Tokyo Sci. Mus. 6:107, 1943.
Tano-cho, Miyazaki, Japan, leg. T. Nukumizu (neotype selected here, TFM-F-2420).
= Phellinus xeranticus (Berk.) Pegler, as already pointed out by Imazeki et al.
(1966). For a detailed description of this species, see Pegler (1964) and Ryvarden &
Johansen (1980). Coloured pictures are given in Imazeki & Hongo (1989) and
Imazeki et al (1990).
conchiformis, Daedaleopsis Imaz., Bull. Tokyo Sci. Mus. 6:77, 1943. Accepted in
the genus.
Sendai, Miyagi, Japan, on Quercus serrata Thunb., May 1907, leg. A. Yasuda, (type in
BPI as type of Trametes tricolor Lloyd, BPI-US0321332).
Basidiocarp sessile; pileus dimidiate, applanate, imbricated, 2-6 cm long, 0.3-0.6 cm
thick, margin thin and acute, deflexed; pileus surface cinnamon to chestnut brown,
glabrous, subzonate, concentrically sulcate or not, radially ridged and partly rough;
tubes pale brown, 1-4 mm deep; pore surface grayish brown, pores mostly regular
and angular, partly elongated, (1-)2-3 /mm, dissepiments thin and becoming lacerate;
context corky, pale brown, 1-2 mm thick.
Hyphal system trimitic, contextual generative hyphae hyaline, thin walled, 2-3 um,
with clamp-connections; contextual skeletal hyphae thick-walled, hyaline to pale
yellow, unbranched to occasionally branched, 3-7.5 um wide in KOH (2.5-5 um wide in
DDDDDODDOD
D
10 um
yon”
Fig. 3. Antrodiella gypsea A) basidiocarps, B) section of basidiocarp, C) cystidia, D)
basidiospores, E) basidia, F) contextual generative hyphae, G) contextual skeletal
hyphae. From the lectotype.
a3
Melzer's reagent); contextual binding hyphae thick-walled, hyaline to pale yellow,
conspicuously branched, up to 5m wide in KOH; tramal hyphae as in context
hyphae; basidia and basidiospores not seen in type material (Fig.2, A-C).
Remarks: This species is closely related to D. confragosa, but has smaller pores and
basidiocarps. Spores are not known from the type nor from other collections, but
other microscopic characteristics are rather similar to those of D. confragosa. Cultural
studies and mating tests are needed to decide whether they are distinct species or
only a form of the same species.
It was first named as Trametes tricolor Lloyd as it was taken as a trametoid form of
Lenzites tricolor. As the combination Daedaleopsis tricolor was preoccupied by
Daedaleopsis tricolor (Bull.:Fr.) Bond. & Sing., the name cited above was proposed
by Imazeki.
dickinsii, Daedalea Yasuda, Bot. Mag. Tokyo 36:127, 1922.
Sendai, Prov. Rikuzen (Miyagi Pref.), leg. A. Yasuda (lectotype, selected here,
TNS-202243, isotype in TFM).
Accepted in the genus.
This is only a partly daedaleoid form of Trametes dickinsii Cke. as pointed out by
Yasuda himself (1923). Ryvarden (1988) placed this species in synonymy with
Daedalea incana (Lév.) Ryv. However, a comparison and a renewed examination of
the type from the Paris herbarium (PC) has shown that D. incana has much smaller
pores and a more brownish context. Thus, we concluded that D. dickensii is a
distinct species.
Trametes dickinsit Cke. (1891) has priority over D. dickinsii Yas. Yasuda (1922),
but Cooke's name cannot be transferred to Daedalea, since the name is preoccupied
by that of Yasuda. His name will then be the valid name as long as the species is
placed in Daedalea. In all other combinations, the basionym must be that of Cooke.
Basidiocarp sessile, annual to perennial, pileus dimidiate, applanate to triquetrous,
up to 20 cm long, 10 cm wide, and 5 cm thick, margin usually dull; pilear surface pale
ochraceous to pinkish buff, inner parts often brownish in old specimens, glabrous,
smooth or with scattered nodulae or irregularly rough, concentrically sulcate or not;
tubes up to 15 mm deep;, pores usually circular, partly elongated to daedaleoid,
1-2/mm, dissepiments thick and smooth; context tough corky, pinkish buff, up to 20
mm thick.
Hyphal system trimitic; contextual generative hyphae hyaline, thin-walled, 1.5-3 um
wide, with clamp-connections; contextual skeletal hyphae hyaline to pale brown,
thick-walled to almost solid, unbranched to occasionally branched, 2.5-4.5 um wide;
contextual binding hyphae moderately branched, up to 4.5 um wide; tramal hyphae as
in context; type sterile, so following descriptions of basidia and basidiospores were
made based on TFM-F-16200; basidia 18-25 x 4.5-6 um, 4-sterigmate, with a basal
clamp; basidiospores cylindrical, hyaline, IKI~ 4-5.5 x 1.8-2.5 um (Fig.2, D-I).
Remarks: The species is very common in temperate areas of Japan, most commonly
on Fagus crenata and Quercus spp. Bakshi (1971) reported the species from
Himalayas (specimens in DD examined), and Hjortstam & Ryvarden (1984),
34
10 um
Fig. 4. A-D: Tyromyces incarnatus A) basidiospores, B) basidia, C) tramal
generative hyphae, D) contextual generative hyphae. A and B from TFM-F-15116; Cs
and D from the lectotype. E-H: Echinodontium japonicum E) basidiospores, F)
cystidia, G) generative hyphae, H) skeletal hyphae. from the type.
35
Ryvarden et al. (1986) reported this from Nepal and China respectively under the
name of D. incana. It is probably widespread in temperate areas of eastern Asia.
There is no doubt that D. dickinsii is closely related to the widespread Daedalea
quercina Fr., separated only by the smaller and more regular pores. Comparative
cultural studies and interfertility tests are desirable to ascertain their
interrelationship.
gypseus, Polystictus Yasuda, Bot. Mag. Tokyo 32:249, 1918.
Sendai, Rikuzen (Miyagi), Japan, 26 May 1912, leg. A. Yasuda ( lectotype, selected
here, TNS-201855, isotype in TFM).
= Antrodiella gypsea (Yasuda) Hattori & Ryvarden comb. nov., basionym as cited
above.
Basidiocarp sessile, effused-reflexed or occasionally fully resupinate, annual, very
light in weight; pilei dimidiate or laterally fused and elongated, imbricated or single,
up to 2 cm long and 1 cm wide in single pilei, up to 3 mm thick, margin usually acute;
pilear surface pubescent, radially sulcate or not, white; tubes white, up to 1.5 mm
deep; pore surface white, pores angular, 5-6/mm; context soft-fibrous, white, up to 2
mm thick.
Hyphal system dimitic; contextual generative hyphae hyaline, thin to slightly thick-
walled, 1.5-2 um wide, with clamp-connections, contextual skeletal hyphae hyaline,
thick walled, almost straight, 2-3 tum wide; tramal generative hyphae hyaline, 1.5-2
uum wide, with clamp-connections; tramal skeletal hyphae hyaline, thick to almost
solid, 2-2.5 um wide; cystidia scattered, acute, thin to slightly thick-walled at the tip,
encrusted at the tip or not; hyphal pegs abundant; basidia ellipsoid, 4-sterigmate, 6-9
x 3-4,.2 tum; basidiospores long ellipsoid, hyaline, IKI, thin walled, 2.2-3 x 1.0-1.5 um
(Fig.3).
Remarks. The dimitic hyphal system with clamped generative hyphae, small spores,
a fleshy consistency, and a white rot, make this a typical member of Antrodiella. It is
common in Japan on coniferous trees including species of Abies, Chamaecyparis and
Cryptomeria.
A. gypsea is similar to Oxyporus cuneatus (Murr.) Aosh. with the same white and
soft-fibrous basidiocarp and host range, but the latter species has larger pores, and
the hyphal characteristics are very different.
hispida, Protodaedalea Imaz., Rev. Mycol. 20:159, 1955.
Oomiya-guti, Mt. Fuji, 4 Aug. 1954, leg. Toki (lectotype, selected here in TNS,
isotype in TFM).
= Accepted in the genus.
For a detailed description of this interesting poroid heterobasidiomycete, see
Bandoni et al. (1981).
incarnatus, Tyromyces Imaz., Bull. Gov. Forest Exp. Sta. Tokyo 67: 31, 1954.
Asakawa, Tokyo, Japan, on rotten wood of Abies firma Sieb. & Zucc., 30 July 1938,
leg. R. Imazeki (lectotype selected here, TNS; isotype in TFM).
= Tyromyces incarnatus Imaz.
Basidiocarp sessile, annual; pileus spatulate, flabelliform to dimidiate, applanate, up
36
to 10 cm long, 1 cm thick, margin thin and acute; upper surface pink, rose to reddish
orange, drying darker, glabrous, smooth to somewhat nodulose; tubes up to 5 mm
deep, pinkish white; pore surface pinkish, pores circular, 3-4/mm, dissepiments thin;
context sappy when fresh, drying rigid, up to 7 mm thick.
Hyphal system monomitic; contextual generative hyphae hyaline, thin to slightly
thick walled and with clamps, 2.5-6 [um wide, tramal generative hyphae hyaline, thin-
or thick-walled to almost solid, 2-5 tum wide; cystidia none. Type sterile and
contaminated, and the following descriptions of basidia and basidiospores were made
based on TFM-F-15116; basidia clavate, 4-sterigmate, with a basal clamp, 12-16 x
4.5-5.5 um; basidiospores hyaline, IKI-, thin walled, 4-5 x 1.5-2.5 um (Fig.4, A-D).
Remarks: The species is remarkable by its reddish basidiocarp and host range
(always on conifers) within the genus Tyromyces. Macroscopically the species
remind about Leptoporus mollis (Pers.:Fr.) Quél. which however has simple septate
hyphae.
japonica, Trametes Yasuda, Bot. Mag. Tokyo 32:356, 1918.
Sumoto, Awaji, Japan, 17 Mar. 1918, leg. J. Matsuzawa. (lectotype, selected here,
BPI -US0320326)
= Perenniporia japonica (Yasuda) Hattori & Ryvarden comb. nov., basionym as
cited above. q :
This is a prior name for Perenniporia fulviseda (Bres.) Dhanda (Basionym: Poria
fulviseda Bres., BPI!). For a detailed description of this characteristic species, see
Keller (1986) as P. fulviseda. Besides central Europe, it is also known from the
Himalayas (Thind & Dhanda 1981), specimen in BPI examined).
japonicum, Echinodontium Imaz. J. Jap. Bot. 11:520, 1935.
Hana-yama, Nara, Japan, 11 May 1933, leg. R. Imazeki, on Quercus glauca Thunb.
(type TNS-200301, isotype in TFM).
= Accepted in the genus.
The species is described in Gross monograph of the genus (Gross 1964) where
clearly points to the close relationship to the East American species E. ballouii
(Banker) Gross. Never the less, to draw the attention to this rare species it is
redescribed here.
Basidiocarp effused-reflexed to almost resupinate, perennial; pileus irregular to
lacking, up to 4 mm wide; pilear surface almost black, crustaceous, glabrous, smooth
to slightly rough; hymenophore spiny, spines sordid white to pale buff, rigid, up to 10
mm long, up to 0.5 mm in diameter, often flattened or forked; context woody hard,
cinnamon, up to 1 mm thick, with black shiny crust next to substrate.
Hyphal system dimitic; contextual generative hyphae thin to slightly thick-walled,
hyaline to pale yellow, simple septate with some scattered clamps, 2-4 um wide;
contextual skeletal hyphae thick walled, pale brown, occasionally encrusted at the
tip, 3-6 um wide; tramal hyphae as in context hyphae; cystidia abundant, thick-walled,
pale brown, apically encrusted, mostly imbedded in the trama, 30-50 x 7.5-10.5 um;
basidia not seen; basidiospores ellipsoid, finely echinulate, hyaline, strongly amyloid
in Melzer's reagent, 5.5-7.5 x 3.2-4.5 um (Fig.4, E-G). See, also Imazeki (1935).
of
Remarks: E. japonicum is a very rare species in Japan and restricted to species of
Quercus.
kanehirae, Polyporus Yasuda, Bot. Mag. Tokyo 35:205, 1921.
Mt.Yura, Taiwan, 3 Apr. 1918, leg. R. Kanehira. (type TNS-201566, isotype in TFM)
= Phellinus kanehirae (Yasuda) Ryv.
This species is apparently related to Phellinus nilgheriensis (Mont.) Ryv. (Basionym
Polyporus nilgheriensis Mont., cotype in K selected here as lectotype). The only
distinction between them is the occurrence of setae which are abundant in P.
kanehirae and extremely sparse in P. nilgheriensis. Thus, it is a possibility that they
represent forms of the same species. For a detailed description of P. kanehirae, see
Ryvarden (1990).
kiyosumiensis, Fomitopsis Imaz. & Sasaki nom. nud. in S. Ito, Mycol. Flora of
Japan Vol.2 No.4, 1955.
= Perenniporia latissima (Bres.) Ryv. as already indicated by Imazeki & Hongo
(1965).
komatsuzakii, Polyporus Yasuda, Bot. Mag. Tokyo 31:329, 1917.
Isozaki, Iyo, Japan, 10 Dec. 1916, leg. M. Komatsuzaki. (lectotype, selected here,
TNS-201420)
=Piptoporus soloniensis (Dub.:Fr.) Pilat.
The type is sterile but spore measurements given by Yasuda are those of P.
soloniensis, and they have the same hyphal system. The context of the type is
extremely fragile and grayish compared with P. soloniensis. For the time being, we
prefer to leave it as a synonym of that species.
kusanoana, Trametes Imaz. Bull. Tokyo Sci. Mus. 6:73, 1943.
Bot. Gdn. Tokyo, Japan, 1901, leg. S. Kusano (type of Daedalea kusanoi, Murr., NY).
= Lenzites vespacea (Pers.) Ryv.
The name cited above was proposed by Imazeki as nomen novum for Daedalea
kusanoi Murr. as the combination "Trametes kusanoi" was preoccupied by Trametes
kusanoi (Murr.) Sacc. & Trott. (basionym Coriolellus kusanoi Murr.).
minutissima, Trametes Yasuda, Bot. Mag. Tokyo 34:29, 1920.
Fujishiro, Ibaraki, Japan, 1 Oct. 1919, leg. Y. Irie. (Neotype selected here,
TNS-202074)
= Perenniporia minutissima (Yasuda) Hattori & Ryvarden comb.nov., basionym as
cited above.
Basidiocarp sessile, single, annual; pileus dimidiate, triquetrous to applanate, 75 x 60
mm in type specimen (up to 10 cm long and 2 cm thick in other specimens in fresh
condition), margin thick and dull; pilear surface reddish brown, almost white near
margin, glabrous, irregularly rough to almost smooth; tubes white, 5 mm deep in type;
pore surface white, pores circular, 3-5/mm; context white, woody hard when dry
(waxy and fragile with abundant semitransparent spots in fresh condition), up to 8
mm thick in type.
Hyphal system di-trimitic; contextual generative hyphae hyaline, thin to slightly
thick-walled, with clamp-connections, 2-3.5 tum wide; contextual skeletal-binding
38
Fig. 5. Perenniporia minutissima A) basidiocarps, B) section of basidiocarp, C)
basidiospores, D) basidia, E) contextual generative hyphae, F) contextual skeletal-
binding hyphae. from the neotype.
39
hyphae hyaline, dextrinoid in mass, thick-walled, 2.5-5 um wide; tramal hyphae
agglutinated; cystidia none; basidia 4-sterigmate, with a basal clamp, 20-30 x 9-12.5
uum; basidiospores ellipsoid, truncate at the apex, thick walled, hyaline, dextrinoid in
Melzer's reagent, 10-13 x 6-7.5 um (Fig.5).
This species is remarkable within the genus Perenniporia by its waxy consistency of
the context in fresh condition. Most collections seen so far have been made on
Symplocos spp. Some specimens were also collected on Pinus densiflora.
Microscopically the species comes close to P. ochroleuca (Berk.) Ryv. which
however has rather small triquetrous perennial basidiocarps with a pale ochraceous
upper side.neo-japonicum, Ganoderma Imaz., Bull. Tokyo Sci. Mus. 1:37, 1939.
Mt.Takao, Tokyo, Japan, Aug. 1935, leg. Y. Kobayasi (type, TNS- 200762).
This species is a member of the Ganoderma lucidum complex. For the time being, we
prefer to keep it separate as Ganoderma neo- japonicum maz.
Basidiocarp laterally or occasionally centrally stipitate, annual; pileus reniform to
dimidiate, applanate, margin thin and deflexed, 3-11 x 4-11 cm; pilear surface almost
black, glabrous, laccate and shiny, radially ridged; tubes dark cinnamon, 0.2-1 cm
deep; pore surface sordid white to cinnamon, pores angular to circular, 4-5/mm;
context cream white near pilear surface, dark cinnamon near tubes, fibrous-corky, up
to 7 mm thick; stipe erect, 9 cm long in type material (up to 27 cm long in other
specimens), stipe surface black and laccate.
Hyphal system di-trimitic; contextual generative hyphae not seen in the type; context
skeletal-binding hyphae pale yellow to brown, rarely to conspicuously branched, up to
7.5 um wide; tramal hyphae as in context; basidia not seen; basidiospores ellipsoid,
truncate at the apex, pale brown, wall two-layered with inter-wall pillars between the
layers, (8.5-)9.5-13 x 6-8 um.
The species is remarkable with its slender long spores, a black shiny basidiocarp,
slender stipe and host range (always on coniferous trees). Coloured pictures are
given in Imazeki & Hongo (1989) and Imazeki et al (1988).
nipponica, Daedaleopsis Imaz., Bull. Tokyo Sci. Mus. 6:78, 1943.
Nikko, Tochigi, Japan, 7 Nov. 1938, leg. R. Imazeki (type, TNS - 208958).
= Daedaleopsis purpurea (Cke.) Imaz. & Aosh. as already indicated by Imazeki et
al. (1966) where also a detailed description can be found.
In Japan, D. purpurea is common on hardwood especially on Prunus spp. The species
is characterized by its regular pores and a multizonate, red to bay pileus surface. The
latter character is shared with D. tricolor and D. styracina (P.Henn. & Shir.) Imaz.
which however have lamellae, and lamellae or large irregular pores respectively.
nipponicus, Polystictus Yasuda, Bot. Mag. Tokyo 30:297, 1916.
Mt.Akagi, Kotsuke, Japan, 28 Sep. 1915, coll. K. Tsunoda (lectotype, selected here,
TNS-201758, isotype in TFM).
= Trametes spp.
The type is thin, flabelliform and sterile and in general in bad condition. No reliable
conclusion can be reached as to its true identity and the name should be dropped from
consideration as a nomen ambigum.
Fig. 6 Trichaptum parvulus A) section of basidiocarp, B) section of hymenium with
cystidia, C) cystidia, D) basidiospores. From the type.
41
orientalis, Polystictus Yasuda, Bot. Mag. Tokyo 32:135, 1918.
Sendai, Japan, 15 Sep. 1912, leg. A. Yasuda (lectotype, selected here, TNS-201876).
= Trametes cfr. lactinea Berk.
Specimens called Trametes orientalis (Yasuda) Imaz. in Japan have often a grayish
and glabrous pileus, different from the white to pale leather coloucsed pileus of T.
lactinea, a species which is fairly widespread in Japan. Cultural studies are
necessaryto verify if this is only a form of T. lactinea.
parvulus, Irpex Yasuda, Bot. Mag. Tokyo 35:254, 1922.
Sendai, Japan, 1 Oct. 1916, leg. A. Yasuda (lectotype, selected here, TNS-203047,
isotype in TFM). (Fig.6).
= Trichaptum parvulus (Yasuda) Hattori & Ryvarden comb. nov. Basionym as cited
above.
Basidiocarp annual, resupinate, adnate but margin slightly lifted in dry condition,
margin narrow to lacking; hymenophore ochraceous to pale brown with a distinct pale
pink hue, irpicoid to hydnoid and crowded with flattened spines, 2-3 per mm, up to 1
mm long, subiculum 0.3 mm thick, dense and cartilaginous as the spines in dry
condition.
Hyphal system dimitic, generative hyphae hyaline, with clamps, 2-4 um wide,
skeletal hyphae dominating in the basidiocarp, hyaline, thick-walled to solid, 3-6 um
wide, hymenial cystidia very abundant, ventricose to cylindrical, thick-walled and
most of them with an apical encrustation, 10-15 x 4-6 lim; present also in the
hymenium are skeletal cystidia arising deep in the central part of the spine as
skeletal hyphae and bending into the hymenium as cystidia, often with apical
encrustation; basidia not seen, spores hyaline, thin-walled, smooth, negative in
Melzer's reagent, ellipsoid, 6-7 x 3 um. Type of rot unknown, on hardwoods, the type
was collected on Lespedezia burgeri Mich. (Fabaceae).
Besides the type, also the following collection was examined: Japan, Chiba
Prefecture, Kiyosumi Forest Sta, Tokyo University Forest. 24. August 1983. L.
Ryvarden 20866 (O and TKB).
Maas Geesteranus (1974:493) has commented upon the status of this species and
pointed out the resemblance to Steccherinum although he felt the structure and shape
of the spines excluded the species from that genus. We feel that the species belongs
in Trichaptum because of the irregular hymenophore, also seen in T. fuscoviolaceus,
the cartilaginous consistency which is so typical for many Trichaptum species and
numerous small ventricose to clavate apically encrusted cystidia, a feature seen in
many Trichaptum species. The spores come close to those of T. byssogenus, but are
slender and basidiocarps of that species normally are poroid and pileate and when
resupinate, are pale umber brown, and have much larger pores which however may
split. its configuration is thus grossly different from the species discussed here.
purpureus, Irpex Yasuda ex Lloyd, Mycol. Writ. 5:715, 1917.
Mt.Iwaya, Iyo, Japan, 19 Oct. 1916, leg. M. Komatsuzaki (lectotype selected here,
BPI -USO325519).
= Australohydnum dregeanum (Berk.) Hjorts. & Ryv. (basionym Corticium
42
Fig. 7. Ganoderma tsunodae A) basidiocarp, B) section of basidiocarp, C)
basidiospores, D) contextual vegetative hyphae. From the lectotype.
43
dregeanum Berk., K!). For a detailed description of this species, see Hjortstam &
Ryvarden (1990).pusilla, Coltricia Imaz. & Ysk. Kobayashi, Trans. Mycol. Soc.
Japan 7:42, 1966.
Kiyomizu, Kyoto, Japan, 18 Aug. 1991, leg. T. Hattori, (neotype, selected here TNS,
isotype in TFM, O).
= Coltriciella pusilla (Imaz. & Ysk. Kobay.) Corner
This species is remarkable by its spatulate and laterally stipitate basidiocarp. For a
detailed description, see Imazeki & Kobayashi (1966). A coloured picture is provided
in Imazeki & Hongo (1989).
It should be noted that the macroscopical characters of this species are identical with
those of the C. dependens (Berk. & Curt.) Murr., which however, has a pendent
basidiocarp. Cultural studies are desirable to ascertain their status.
Morphologically we are convinced that they represent two independent species as no
intermediate forms have ever been collected. Further, the upper surface of C.
dependens is velutinous and usually with hairs on stipe in fresh condition, while it is
glabrous and shiny in C. pusilla. The context in C. dependens is fibrous-spongy and
very light in weight in dried condition while it is leathery in C. pusilla as for example
in Coltricia cinnamomea.
sciurinus, Inonotus Imaz., Bull. Tokyo Sci. Mus. 6:106, 1943.
Nikko, Totigi, Japan, 7 Nov. 1938, leg. R. Imazeki (type TNS- 206965, isotype in
TFM).
= Inonotus flavidus (Berk.) Ryv. (basionym Polyporus flavidus Berk., K!).
The species is characterized by a thin black line in the duplex context, a thick
tomentum, small cylindrical spores and presence of scattered hymenial setae.
Besides Japan, it is widespread in the Himalayas.
sendaiensis, Polyporus Yasuda, Bot. Mag. Tokyo 37:101. 1923. Sendai, Japan, 21
Sep. 1913, leg. A. Yasuda (type TNS, isotype in TFM).
= Pyrroderma sendaiense (Yasuda) Imazeki,
For a detailed description, see Imazeki (1966) and Ryvarden (1991) with a key to
accepted species in the genus.
sendaiensis, Trametes Yasuda, Bot. Mag. Tokyo 36:21, 1922.
Sendai, Rikuzen (Miyagi), Japan, 11 Aug. 1921, leg. A. Yasuda (lectotype, selected
here, TNS, isotype in TFM).
= Datronia stereoides (Fr.) Ryv.
symploci, Trametes Yasuda, Bot. Mag. Tokyo 37:84, 1923.
Bessho, Yahara, Prov. Hitachi (Ibaraki pref.) Japan, 17 Oct. 1918, leg. Y. Irie
(lectotype, selected here, TNS)
As already pointed out by Yasuda himself in the original description (1923), this
name is a later synonym of Perenniporia minutissima (Yasuda) Hatt. & Ryv.
subumbraculum, Ganoderma Imaz. Bull. Tokyo Sci. Mus. 1:38, 1939.
Hane-yama, Koti, Japan, Sep. 1908, leg. K. Ogawa (holotype, TNS- 206977). The
species is apparently a member. of the Ganoderma lucidum complex. This complex is
in a taxonomic chaotic state, and thus, for the time being, we leave the species under
A4
Imazeki's name.
The species is known only from the type. The most striking character are the
centrally stipitate basidiocarp, the light- coloured context with 3 layers, and the light
weight in dried state. Fig. 7.
tabacinoides, Irpex Yasuda, Bot. Mag. Tokyo 34:96, 1920.
Prov. Harima, Kashima-mura, Ibo-gori, 2 Aug. 1918, on Pasania cuspidata (Thunb.)
Oerst. (= Castanopsis cuspidata (Thunb.) Schottky), leg. U. Ouye (Lectotype
selected by Ryvarden (1982), TNS -204262).
= Hydnochaete tabacinoides (Yasuda) Imaz.
For a detailed description, see Ryvarden (1982).
tsunodae, Polyporus Yasuda ex Lloyd, Mycol. Writ. 5:792.
Ikeda, Kozuke, Japan, 8 July 1917, leg. K. Tsunoda, (lectotype selected here,
BPI -US0307263).
= Ganoderma tsunodae (Yasuda ex Lloyd) Trott.
Basidiocarp sessile, single, annual; pileus dimidiate, flabelliform to spatulate,
applanate, 3-12 cm long, 3-6.5 cm wide, 1.5-3 cm thick; pilear surface dark cinnamon
to ochraceous, coarse, glabrous; tubes sordid white, up to 15 mm deep; pore surface
white, pores round to angular, 4-5/mm; context white, fibrous-corky and very rigid
when dry (watery-fibrous when fresh).
Hyphal system di-trimitic; contextual generative hyphae not seen in the type;
contextual skeletal-binding hyphae hyaline, thick-walled to almost solid, occasionally
to conspicuously branched; tramal hyphae as in context; basidia not seen;
basidiospores ellipsoid to truncate, pale yellow, wall two layered with interwall
pillars, 20-24 x 14-16.5 um (Fig.8).
This species is remarkable in Ganodermataceae with its white, watery context and
large and yellow spores. Macroscopically it reminds remarkably about G. colossus
(Fr.) Baker, which however has smaller spores (14-19 x 8-12 [1m).
Imazeki (1939) proposed Trachyderma for G. tsunodae. However, the generic name
is illegitimate as a homonym of Trachyderma Norm. 1853 as pointed out by Ryvarden
(1991). On the other hand, G. colossus is the type species of Thomophagus Murr.
1905. Thus, should it ever be desirable to place these two species in a genus of its
own due to their loose and soft consistency, the latter name is available.
yamanoi, Cryptoderma Imaz., Forsch. Geb. Pflanzenkrankh. 4:176, 1951.
Iburi, Lakeside of Sikotu, on Picea jezoensis (Sieb. & Zucc.) Carrié, 10 Sep. 1948,
leg. R. Imazeki (type, TFM-F-1134, isotype in TNS).
= Phellinus pini (Fr.) Quél., as already pointed out by Imazeki & Hongo (1989).
Cerny (1985) concluded that Phellinus vorax (Hark.) Cerny is distinct from P. pini
with a different ecological niche and morphology. If P. vorax is accepted as an
independent species, C. yamanoi is a synonym of this species.
ACKNOWLEDGMENTS
We are most grateful to Professor R.L. Gilbertson for a critical review of the
manuscript. Dr. Y. Doi, National Science Museum, kindly deposited some type
45
materials to Herbarium of For. & For. Prod. Res. Inst. (TFM). The curators of
following herbaria are greatly appreciated for the loan of type materials and other
specimens: National Science Museum (TNS), U. S. National Fungus Collections
(BPI), New York Botanic Garden (NY), Royal Botanic Gardens (K), Muséum
National d'Histoire Naturelle (PC), Forest Research Institute, India (DD). T. H. is
also grateful for Dr. R. L. Gilbertson, Univ. Arizona, for the loan of a recent collection
of Melanoporia nigra.
REFERENCES
Aoshima, K. & Kobayashi, T. 1983: Wood-rotting fungi found in living Populus and
Salix in Japan. Trans. 94th Mtgs. Jpn. For. Soc.: 545-546 (in Japanese).
Bakshi, B. K. 1971: Indian Polyporaceae, 244 pp, Indian Council of Agricultural
Research, New Delhi.
Bandoni, R., Oberwinkler, F. & Wells, K. 1982: On the poroid genera of the
Tremellaceae. Can. J. Bot. 60:998-1003.
Cerny, A. 1985: Taxonomic study in the Phellinus pini complex. Ceska Mykol.
39:71-84.
Furukawa, H., Abe, Y. & Neda, H. 1983: List of fungi of Mt. Fujii. Trans. Mycol.
Soc. Japan 24:235-245.
Gilbertson, R. L. & Ryvarden, L. 1986 & 1987: North American Polypores Vol. 1 &
2, 885 pp, Fungiflora, Oslo.
Gross, H. L. 1964: The Echinodontiaceae. Mycopath. Mycol. Appl. 24:1-26.
Hjortstam, K. & Ryvarden, L. 1984: Some new and noteworthy Basidiomycetes
Aphyllophorales) from Nepal, Mycotaxon 20:133-151.
Imazeki, R. 1935: Studies on Echinodontium Ellis et Everhart, J. Jap. Bot.
11:514-521 in Japanese with English descriptions).
Imazeki, R. 1943: The genus Grifola S.F.Gray. Polyporaceae of Eastern Asia I, J.
Jap. Bot. 19:381-390 (in Japanese).
Imazeki, R. 1966: The genus Pyrrhoderma Imazeki, Trans. Mycol. Soc. Japan 7:3-11.
Imazeki, R. & Hongo, T. 1965: Colored illustrations of fungi of Japan, Vol.2, 149 pp,
Hoikusha, Osaka (in Japanese).
Imazeki, R. & Hongo, T. 1989: Colored illustrations of mushrooms of Japan, Vol. 2,
315 pp, Hoikusha, Osaka (in Japanese).
Imazeki, R. & Kobayashi, Y. 1966: Notes on the genus Coltricia S.F.Gray, Trans.
Mycol. Soc. Japan 7:42-44.
Imazeki, R., Kobayasi, Y. & Aoshima, K. 1966: Fungi, in The flora of eastern
Himalaya, pp 611-626, Univ. Tokyo, Tokyo.
Imazeki, R., Otani, Y.,& Hongo, T. 1988: Fungi of Japan, 623 pp, Yamakei, Tokyo
(in Japanese).
Ito, S. 1955: Mycological Flora of Japan Vol.2, No.4 450 pp, Yokendo, Tokyo (in
Japanese).
Keller, J. 1986: Ultrastructure des Parois Sporiques de Quelques Aphyllophorales,
Mycol. Helvetica 2:1 -34.
46
Maas Geesteranus, R.A. 1974: Studies in the genera Irpex and Steccherinum.
Persoonia 7:443-581.
Pegler, D. N. 1964: A survey of the genus Jnonotus (Polyporaceae, Trans. Brit.
Mycol. Soc. 47:175-195.
Ryvarden, L. 1982: The genus Hydnochaete Bres. (Hymenochaetaceae, Mycotaxon
15:425-447.
Ryvarden, L. 1988: Type studies in the Polyporaceae 19. Species described by M. C.
Cooke. Mycotaxon 33:303-327.
Ryvarden, L. 1990: Type studies in the Polyporaceae 22. Species described by
C.G Lloyd in Polyporus , Mycotaxon 38:83-102.
Ryvarden, L. 1991: Genera of polypores, nomenclature and taxonomy, Syn. Fung.
5:1-363, Fungiflora, Oslo.
Ryvarden, L. & Gilbertson, R. L. 1993: European Polypores, part 1. Synop. Fung.
6:1-387. Fungiflora, Oslo.
Ryvarden, L. & I. Johansen 1980: A preliminary polypore flora of East Africa, 636
pp, Fungiflora, Oslo.
Ryvarden, L., Xu Liang-Wang, & Zhao Ji-Ding 1984: A note of the Polyporaceae in
the Chang Bai Shan Forest Reserve in Northern China, Acta Mycol. Sin. 5:226-234.
Thind, K.S. & Dhanda, R. S. 1980: The Polyporaceae of India XIII. Indian
Phytopatol. 33:380-387.
Yasuda, A. 1923: Zwei neue Arten von Trametes, Bot. Mag. Tokyo 37:83 -85.
MY COTAXON
Volume L, pp. 47-59 January-March 1994
THE SPECIES OF THE GENUS RINODINA (LICHENIZED
ASCOMYCETES, PHYSCIACEAE) CONTAINING PANNARIN IN
EURASIA WITH A SPECIAL NOTE ON THE TAXONOMY OF
RINODINA GRANULANS
Mireia GIRALT’, Helmut MAYRHOFER™ and Walter OBERMAYER~
“Department of Plant Biology, University of Barcelona, Diagonal 645,
E-08071 BARCELONA, Spain.
Institut far Botanik, Karl-Franzens-Universitat Graz, Holteigasse 6,
A-8010 GRAZ, Austria
ABSTRACT: Six species of the genus Rinodina with thalli and/or
discs PD+ orange due to the presence of pannarin are hitherto
recognized in Eurasia. One is saxicolous, R. santorinensis, and
five are corticolous, R. dalmatica, R. efflorescens, R. excrescens,
R. granulans and RA. pruinella. Notes on the taxonomy of A.
granulans are provided. Descriptions and illustrations of this
species and the also poorly understood A. excrescens are
included. A key to the treated species is given. Other known
species containing pannarin are briefly discussed.
INTRODUCTION
The treated species are specially characterized by the content of pannarin. This
lichen substance is usually concentrated in the thallus, in the thalline exciple
and/or in the epihymenium which appear entirely interspersed with irregularly
shaped and sized crystals. These crystals are clearly visible under polarizing
light and dissolve after treatment with PD forming reddish-orange acicular
crystals.
Depending on several characters the treated species can be included in four
different groups:
Rinodina efflorescens, R. excrescens and R. granulans are very closely related.
They have in common the following characters: (1) pannarin as a principle
lichen substance concentrated in the thallus, the thalline exciple and the
epihymenium; (2) a well developed thallus, composed of scattered to
48
contiguous areolae, appearing as minute flattened squamules which dissolve
into soredia or blastidia; (3) an epihymenium with a granular and PD+ orange
epipsamma (pannarin); (4) thalline exciple reacting 1+ blue (5) asci of the
Lecanora-type (HONEGGER 1978) and (6) ascospores of the Physcia-type with
tendencies to the Milvina-type.
Rinodina dalmatica is separated from the first group only because of the
Pachysporaria-type ascospores.
Rinodina pruinella differs from the species cited above in the following
characters: (1) pannarin located only in the epihymenium. Atranorin constitutes
the principle lichen substance; (2) thallus not composed of areolae and not
developing vegetative propagules; (3) thalline exciple I- and (4) ascospores of
the Dirinaria-type.
Lastly, Rinodina santorinensis s. \at. is distinguished by: (1) pannarin present
only in the thallus and the thalline margin; (2) epihymenium PD-, without
epipsamma and (8) habitat saxicolous and usually parasitic.
According to SHEARD (in lit.), the literature checked and our own investigations,
other species hitherto described possessing pannarin but not present in the
study area are: the saxicolous Rinodina murrayii H. MAYRHOFER, from
Australasia, and the corticolous R. adirondackii H. MAGN., R. granuligera H.
MAGN., AR. marysvillensis H. MAGN. and RA. thujae (H. MAGN.) SHEARD, all from
North America. Complete descriptions of these species are given by
MAYRHOFER (1983) and MAGNUSSON (1932, 1947b and 1953).
R. murrayii belongs to the same group that R. excrescens, R. granulans and R.
efflorescens, but the thallus does not develop vegetative diaspores and is
saxicolous. AR. adirondackii, like R. dalmatica, is separated by its
Pachysporaria-ascospores, but its thallus does not develop vegetative
propagules. R. granuligera and R. marysvillensis are closely related to R.
pruinella but their ascospores belong to the Physcia-type.
We cannot exclude the possibility that other pannarin containing species may
exist because the PD reaction was often not checked by other authors.
The terminology employed for the vegetative propagules follows that of POELT
(1980), HAWKSWORTH et al. (1983) and Fox & PURVis (1992), for the ascospores
MAYRHOFER (1982) and SCHEIDEGGER (1993) and for the asci HONEGGER (1978).
For the identification of lichen substances the standardized methods for thin
layer chromatography (TLC) were used (e.g. CULBERSON & AMMANN 1979).
KEY TO THE SPECIES
ta Saxicolous. 2
1b Corticolous or lignicolous. 3
2a Epihymenium PD-. Ascospores Pachysporaria-type with tendencies to
Physcia-type, 14-22 x 7-14 um. Thallus parasitic or seldom autotrophic.
Macaronesia and mediterranean region. R. santorinensis
2b Epihymenium PD+ orange. Ascospores Physcia-type, 16-22 x 10-13 um.
Thallus always autotrophic. Australasia. (R. murrayii )
49
3a _ Thallus lacking of vegetative propagules, PD+ faint yellow. Ascospores
Dirinaria-type, 15-25 x 7-13 pum. Maritime, southem mediterranean-
atlantic. R. pruinella
3b Thallus sorediate or blastidiate, PD+ orange. Ascospores different. 4
4a _ Thallus sorediate. Soralia discrete, never forming a continuous leprose
crust. Ascospores Physcia-type, 15-20 x 7-10 um. Oceanic, boreal and
suboreal. R. efflorescens
4b Thallus blastidiate. Blastidia forming a more or less continuous
granulose-isidiose or leprose crust. 5
5a_Blastidia large, up to 60-80(-100) um diam., forming a granulose-isidiose
crust (appearing subSquamulose). Ascospores Physciatype, 14-21 x 7,5-
11 um. Boreal, subboreal and mediterranean R. excrescens
5b Blastidia small, up to 30 um diam, forming a continuous leprose crust. 6
6a Ascospores Pachysporaria-type, when young with polygonal lumina, 17-
26 x 8-13 um. Maritime, mediterranean-atlantic. AR. dalmatica
6b Ascospores Physcia-type, when young never with polygonal lumina, 18-
25 x 10-14 um. Siberia. R. granulans
THE SPECIES
1) RINODINA DALMATICA ZAHLBR.
Osterr. Bot. Z. 51: 348 (1901); BOULY DE LESDAIN (1909: 170); HARMAND (1913:
902); MAGNUSSON (1947a: 313); GIRALT et al. (in prep.).
Type: Croatia: Dalmatia, "in peninsula Lapad prope Ragusa", 100 m, on Pinus
halepensis, J. BAUMGARTNER (GZU, W, WU-isotypes).
Exs.: ZAHLBRUCKNER: Lich. rar. exs. 39 (GZU, W, WU).
Rinodina dalmatica is distinguished by the entirely blastidiate thallus forming a
continuous leprose crust and the Pachysporaria-type ascospores, when young
with typical polygonal lumina. It is unique amongst Rinodina species in having
the latter feature.
Selected specimens examined: CROATIA: Dalmatia: Ragusa, Mte. Petka,
150 m, on Pinus halepensis, A. LATZEL (W). - GREECE: Korfu: Kerkyra, NW
Ipsos, on Olea, 16-17.8.1970, J. POELT (GZU). - Peloponnes: Elis, Olympia, on
Pinus halepensis, 14.4.1971, J. POELT (GZU). - ITALY: Latium: Roma, Tenuta
di Caccia di Castel Porziano, SW Roma, 0-20 m, 8.5.1986, J. POELT (GZU). -
PORTUGAL: Algarve: Sierra de Monchique, Caldas, Vale do Paraiso, on Olea
europaea, 24.2.1946, C.N. TAVARES & L. SOBRINO (LISU). - Estremadura:
Setubal, Mata do Reboredo, on Pinus halepensis, 14.5.1944, C.N. TAVARES
(LISU).
50
2) RINODINA EFFLORESCENS MALME
Svensk Bot. Tidskr. 21: 251 (1927); MAGNUSSON (1947a: 229); HARRIS (1977:
129, as Rinodina sp. 1, according to SHEARD in lit.); COPPINS & JAMES (1979:
175); WITTMANN & TURK (1987: 394); DIEDERICH (1989: 204); ETAYO (1992: 192);
Fox & PURVIS (1992: 548); TONSBERG (1992: 286); WIRTH (1990: 328); WONG &
BRODO (1990: 364, 1992: 70); GIRALT et al. (1993: in press.).
Type: Sweden: Vastergotland, Habo, St. Karr, on old Fagus in a shady place,
1923, G.O. MALME (S - not seen).
Syn.: Rinodina hueiana (HARM.) OLIV., Bull. Acad. Internat. Geogr. Bot. 15: 211
(1905). - Lecanora hueiana HARM., Bull. Soc. Sci. Nancy, ser. 2, 15: 195
(1898); HARMAND (1913: 883). - non Rinodina hueana VAIN., Hedwigia 37: 38
(1898). - Type: France: Vosges, "en montant au Ballon d'Alsace depuis Saint
Maurice, sur un Bouleau, parasite sur le thalle vieux du Parmelia saxatilis",
HARMAND (ANGUC - not seen).
Discussion: The incorrectly spelled epithet hueiana [see recommendation
73C(c) of the |.C.B.N.] is only an orthographic variant of hueana and the
combination Rinodina hueiana proposed by OLIVIER (loc. cit.) is a latter
homonym of Rinodina hueana VAIN. DIEDERICH (1989) already listed R. hueiana
as a synonym of R. efflorescens. WIRTH (1990) placed A. efflorescens in the
synonymy of AR. hueiana without further comments. Because of the mentioned
criteria R. efflorescens is the correct name of this species.
Exs.: Lich. sel. exs. Upsalienses 71 (GZU).
The discrete soralia are diagnostic of R. efflorescens.
Selected specimens examined: AUSTRIA: Steiermark: Nordliche Kalkalpen,
ca. 3 km W of GroBreifling, ca. 640 m, on Acer sp., 26.10.1990, J. POELT, J.
HAFELLNER & E. LOPEZ DE SILANES (GZU). - BRITISH ISLES: Isle of Wight:
Bortheood Copse, on Quercus, November 1977, F. Rose (BM). - Main Argyll:
Seil, Ballachuan, Port Mor, on Sorbus aria, 5.8.1980, P.W. JAMES (BM). - East
Sussex: Eridge Old Park, on old oak, June 1968, F. Rose & P.W. JAMES (BM).
- FRANCE: South-Voges: Steinbach, Cernay, ca. 700 m, on Quercus petraea,
27.10.1968, V. WIRTH (STU). - GERMANY: Warttemberg: Neckar, Leonberg,
Warmbronn, Stdckach, 440 m, 16.10.1988, V. WIRTH (STU). Baden,
Sudschwarzwald, Schdnau, 600 m, 30.9.1971, V. WIRTH (STU). - SPAIN:
Navarra: Belabarce, 1100 m, on Fagus sylvatica, 25.7.1987, J. ETAYO 3313
(Etayo).
3) RINODINA EXCRESCENS VAIN.
Ann. Acad. Sci. Fenn., Ser. A, 27: 84 (1928); MAGNUSSON (1947a: 237); GIRALT
et al. (1993: 711).
Type: Siberia :"Sibiria Occidentali, Konda, ad lignum putridum in pineto prope
Leunsk", 1880, E. VAINIO (TUR-V 08798-holotype).
51
Thallus (Figs. 1c; 2a,b) crustaceous composed of scattered to contiguous
areolae. Areolae light grey to grey brown, plane to bullate, 0,1-0,5 mm diam.,
with the appearance of minute sublobate squamules, becoming more or less
confluent, mostly developing ascending blastidia. Blastidia large, 60-100 um
diam., scattered or confluent and then forming a continuous granulose-isidiose
crust, concolorous with areolae. Apothecia (Figs. 1c; 2a) rare, sessile, scattered
or contiguous, up to 1 mm. Thalline margin concolorous with thallus, at first
thick, entire and prominent, becoming thinner and flexuose, often partially
excluded. Disc plane, rarely convex, brown, slightly pruinose. Proper margin
often visible as a ring within the thalline margin. Excipulum thallinum up to 80
pum laterally, expanded to 100-110 zm below, I+ blue, inspersed with crystals of
pannarin. Cortex cellular, 15-20(-40) um laterally, expanded to 40-60 um
below, cells thin-walled. Excipulum proprium 10-15 ym laterally, expanded to
25-40 ym above. Epihymenium reddish-brown, with a granular and PD+ orange
epipsamma. Hymenium (60-)80-100 um high. Hypothecium colourless 40-110
jum deep. Paraphyses ca. 1,5-2 um wide, apices capitate, 2,5-3,5 um wide.
Asci 8-spored, Lecanora-type. Ascospores (Figs. 2c-e) Physcia-type, (15-)17-
19(-21) x (7,5-)9-10(-11,5) zm, smooth (Someones when overmature minutely
warted), constricted at the septum, with a well developed torus (Fig. 2e).
Spermatia not seen.
Chemistry. Thallus K+ yellow and PD+ orange; pannarin and atranorin by TLC.
DISCUSSION: This species is characterized by the thallus composed of
discrete to contiguous areolae which partially develop ascending blastidia
(isidia-like) and form a more or less continuous granulose-isidiose (appearing
subsquamulose) crust (Fig. 2a-b). The large blastidia, up to 100 um diam.,
make this species morphologically distinct from its relatives.
Rinodina thujae (H. MAGN.) SHEARD, from North America, is closely related if
not conspecific to R. excrescens. The holotype differs from the investigated
specimens of A. excrescens exclusively in the areolae which do not build
structures resembling blastidia (compare Fig. 1d and Fig. 2a-b). Another
sample of RAR. thujae studied (WETMORE 32864) is somewhat intermediate
between both species, with some areolae developing blastidia-like structures
(=ascending and sublobate margins). After the study of this sample, and
without having seen the holotype, this species was mentioned in GIRALT et al.
(1993) as a possible synonym of R. excrescens. The study of further North
American material of R. thujae is necessary in order to solve its taxonomical
position definitively.
HABITAT AND DISTRIBUTION: Ainodina excrescens has hitherto been
reported only from the type locality in West Siberia (VAINIO 1928, MAGNUSSON
1947a) and from Austria (GIRALT et al. 1993). According to SHEARD (in lit.) the
record from North America reported by BRODO et al. (1987) and also cited in
EGAN (1987) refers to another, undescribed Rinodina species.
Additional specimens examined: AUSTRIA: Steiermark: Gurktaler Alpen,
Grebenzen, 6 km NW Neumarkt, 1 km NW Oberdorf, Dirnberger Hochmoor,
ca. 980 m. on Juniperus communis, 20.1.1988, W. OBERMAYER 2587 (GZU,
Obermayer). Weststeirisches Hugelland, zwischen Deutschlandsberg und
oe
Fig. 1. Type specimens of Rinodina granulans (a,b), R. excrescens (c) and R.
thujae (d); a. R. granulans (white soredia) growing together with R. archaea
agg. (3 apothecia). b. apothecium (arrow) of R. granulans. scale = 500 pm.
Fig. 2. Rinodina excrescens (OBERMAYER 2587). a. apothecia. b. blastidiate
areoles. scale = 500 pm. c-e. ascospore ontogeny (Physcia-type ascospores).
scale = 10 um.
54
Schwanberg, ca. 390 m, on Quercus robur, J. HAFELLNER 23250 & M.E. LoPEz
de SILANES (GZU, Hafellner). - CROATIA, Insel Mijet, S Veliko Jezero, 0-30 m,
on Pinus halepensis, 15.04.1979, O. BREUSS 954 (BreuB).
4) RINODINA GRANULANS VAN.
Ann. Acad. Sci. Fenn., Ser. A, 27: 83 (1928)
Syn.: A. sibirica var. granulans (VAINIO) H. MAGN., Acta Horti Gothob. 17: 272
(1947a). - Type: Siberia: "Sibiria Occidentali, Konda, in saepimento ligneo
loco umbroso ad Tumynvatsk", 1880, E. VAINIO (TUR-V-08799 - holotype)
Thallus (Figs. 1a-b) crustaceous, composed of scattered to contiguous areolae.
Areolae whitish, whitish-grey, whitish-green or brownish, matt, plane or slightly
convex (0,1-)0,2-0,3(0,5) mm diam., usually becoming confluent, with the
appearance of minute squamules, dissolving completely into small blastidia.
Photobiont trebouxioid, cells 7-12(-15) mm diam. Blastidia small, 15-30(-50)
uum diam., confluent, forming a continuous leprose crust, whitish (in the
holotype) or with a +dark brown tinge (in the other specimens examined),
budding from the margins and surfaces of the areolae. Apothecia very rare,
sessile or innate in a dense layer of blastidia, constricted at the base, up to 0,3
mm diam. (all of them are very young). Thalline margin persistent, smooth,
concolorous with the areolae. Disc plane, reddish-brown. Excipulum thallinum
80-90 pm laterally, expanded to 130 ym below, |I+ blue, inspersed with crystals
of pannarin. Cortex cellular, 10-15 um laterally, 30-50 um at the base, cells
thin-walled. Excipulum proprium indistinct laterally, expanded to 30-50 um
above. Epihymenium reddish-brown, with a granular and PD+ orange
epipsamma. Hymenium 70-80 pm tall. Hypothecium colourless, + 100 um
deep. Paraphyses 1,3-2 um wide, apices capitate, 2,5-4 um. Asci Lecanora-
type. Ascospores Physcia-type with tendencies to Milvina-type, constricted at
the septum, smooth, with a well developed torus, 18-25 x 10-14 um. Spermatia
not seen.
Chemistry. Thallus K+ faint yellow and PD+ orange; pannarin by TLC.
DISCUSSION: In the sample of the holotype of A. granulans there are two
Rinodina species growing together (Fig. 1a). One species has a thallus
consisting of discrete to contiguous areolae dissolving completely into small
blastidia and forming a continuous leprose crust. This thallus contains pannarin
(PD+ orange) and includes only very few apothecia, mostly very young and
hidden between the blastidia. The excipulum thallinum is I+ blue and entirely
inspersed with crystals PD+ orange (pannarin); the epihymenium is covered
with a granular epipsamma, not dissolving in K and also PD+ orange; and the
ascospores are Physcia-type, with tendencies to Milvina-type, with walls
irregularly thickened.
The second species has a thin, smooth and continuous thallus without positive
reactions. The apothecia are abundant and very well developed. The
excipulum thallinum is l- and without any crystals; the epihymenium is covered
with a coarsely granular epipsamma dissolving in K and PD-; and the
55
ascospores are Physconia-type, 18-21 x 85-10 um, with uniform walls
(SCHEIDEGGER 1993). This species belongs to the Rinodina archaea group.
VAINIO's original description includes characters of both species. Whereas the
description of the thallus is based on the first species: "Thallus verruculoso-
rugulosus, ....... verruculis dispersis aut contiguous, p. p. soredioso-
fatiscentibus, farinosam confluentibus ...... ", the description of the apothecia
fits with the second one: "Apothecia in partibus minus sorediosis thalli sat
crebre evoluta..... Margine tenui, integro.... Excipulum. ..... , jodo non
reagens..... Sporae long 0,017-0,021, crass 0,09-0,011 mm, membrana sat
aequaliter incrassata......".
According to the I|.C.B.N. (art. 9.2, rec. 7B) we choose the species
characterized by an entirely blastidiate and PD+ orange thallus and Physcia-
type ascospores, as the lectotype of A. granulans.
Rinodina granulans is characterized by its small blastidia (soredia-like) forming
a continuous, +brownish leprose crust and its Physcia-type ascospores grading
into the Milvina-type. Apothecia were observed only in the type material (Fig.
1b).
HABITAT AND DISTRIBUTION: Rinodina granulans seems to be a widely
distributed and quite common species in Siberia where it grows mostly on
lignum and on small dry twigs, more seldom on smooth bark, associated with
other lignicolous species such as Rinodina archaea agg., Candelariella vitellina,
Lecanora symmicta and L. varia. Other associated species are: Melanelia
olivacea coll., M. exasperatula, Parmelia sulcata, Xylographa_parallela,
Cyphelium tigillare, Hypocenomyce scalaris, Calicium salicinum and Caloplaca
cerina.
Additional specimens examined: SIBERIA: Jenisejsk, Nasimova, on lignum,
28.6.1876, M. BRENNER (S). Jenisejsk, near the city of Jenisejsk, on lignum,
21.6.1876, M. BRENNER (S). Jenisejsk, Verst, N of Jenisejsk, on old cortex of
Prunus padus, 26.6.1876, M. BRENNER (S). Jenisejsk, Novo Sjolovskoje, on
lignum, 26.9.1876, M. BRENNER (S). Jenisejsk, Troitskij Klosterdorf, on lignum,
13.7.1876, MM. BRENNER (S). Jenisejsk, Potkamina Tanguska, on lignum,
28.9.1876, M. BRENNER (S). Jenisejsk, Asinovo, on dry twigs, 4.7.1876, M.
BRENNER (S). Jenisejsk, Vorogova. on lignum, 30.9.1876, M. BRENNER (S).
Tobolsk, Kalimski, on lignum, dry twigs and Betula sp., 31.5.1876, M. BRENNER
(S). Tomsk, Timskaja, on dry twigs, 1.6.1876, M. BRENNER (S).
5) RINODINA PRUINELLA BAGL.
Nuovo Giorn. Bot. Ital. 11: 79 (1879); complete information about this species
is given by GIRALT & MAYRHOFER (1994).
Type: Italy: Sardinia, Giorgino iuxta Cagliari, on Ficus carica CANEPA (MOD-
holotype).
Exs.: SAMPAIO: Lich. de Portugal 192 (M, UPS).
56
Pannarin located only in the epihymenium, together with the thallus lacking of
vegetative diaspores and K+ yellow (atranorin) and the Dirinaria-type
ascospores, are diagnostic for this species.
Among the species containing pannarin, R. pruinella and the North American R.
granuligera and R. marysvillensis are unique in having this substance
concentrated only in the epihymenium rather than in the thalline and other
apothecial tissues. This feature easily separates these taxa from their relatives.
R. granuligera and R. marysvillensis cannot be mistaken for R. pruinella
because of their Physcia-type ascospores (see below).
The examined specimens are listed in GIRALT & MAYRHOFER (1994)
6) RINODINA SANTORINENSIS J. STEINER s. lat.
Verh. Zool.-Bot. Ges., Wien 69: 55 (1919); a complete information about this
species is given by MAYRHOFER et al. (1993).
Type: Greece: Santorin Island, between Thira and Pyrgos, 4.1911, R.
WETTSTEIN (W-lectotype).
Exs.: FOLLMANN: Lich. sel. exs. 219 (B, GZU, H, LD, W, as A. confragosa). -
Plantae Graecenses, Lich. 504 (GZU). - VEZDA: Lich. rar. exs. 39 (ANUC, BM,
DUKE, GZU, H, HO, M, PRM, STU, TSB, UPS, VBI, Kalb, Lumbsch and
Vezda).
Rinodina santorinensis and the Australasian R. murrayii are the only saxicolous
species hitherto known containing pannarin. For this reason the latter is also
mentioned in the key. The PD- and I- reactions at the epihymenium and
excipulum level, respectively, distinguish R. santorinensis not only from R.
murrayii but also from the other treated species. On the other hand, the
epihymenium lacking pannarin and the Pachysporaria-type ascospores show
that R. santorinensis is more closely related to R. beccariana and to the
corticolous R. roboris (MAYRHOFER et al. 1993) than to the taxa treated in this
contribution.
The specimens examined are listed in MAYRHOFER et al. (1993).
OTHER RINODINA SPECIES CONTAINING PANNARIN NOT PRESENT IN
THE STUDY AREA
Rinodina adirondackii H. MAGN., Bot. Not.: 48 (1947b). - Type: U.S.A.: New
York, Adirondack Mountains, Chapel Pond, near St. Huberts, 1600ft, on cedar
in gully, 1933, J.L. Lowe (UPS-holotype, not seen).
This species is distinguished by the thallus lacking vegetative propagules and
the large Pachysporaria-type ascospores, 21-35 x 15-21 gum. Ainodina
dalmatica and R. santorinensis possess the same ascospore-type but in both
species those are smaller. Furthermore R. dalmatica has a blastidiate thallus
and A. santorinensis is saxicolous.
SA
Distribution: Limited to America's northeastern states, Ontario and Quebec
(SHEARD in lit). No specimen belonging to this species has been investigated.
All information was provided by Dr.Sheard (Saskatoon).
Rinodina granuligera H. MaGun., Bot. Not.: 35 (1947b). - Type: U.S.A.:
Florida, Sanford, on trees, 1909, RAPP (UPS-holotype, not seen).
Pannarin located only in the epihymenium, together with the thallus lacking
vegetative diaspores, K+ yellow (atranorin) and the Physcia-type ascospores of
15-18 x 6,5-8,5 um, distinguish this species from its relatives (see also A.
pruinella).
Distribution: Southern and Eastern North America (SHEARD in lit.).
Specimens examined: U.S.A.: Louisiana: Baton Rouge, N of Terrebonne-
Lafourche parish line on road 309, west of Thibodaux, on Salix, 11.10.1980,
S.C. TUCKER (GZU). 4,7 miles n-NE of Chipola, on gravel road (Parrish road
1044), hardwood forest in ravine, 8.3.1973, S.C. TUCKER 11101B (Sheard).
West Side of lake Bistineau, on dirt road parallel to lake edge, immature mixed
Pinus sp.- Quercus ilex forest, 26.5.1973, S.C. TUCKER (Sheard).
Rinodina marysvillensis H. MAGN., Ann. Cryptog. Exot. 5: 31 (1932). - Type:
U.S.A.: Washington, Marysville, on Salix bark, 1927, GRANT (UPS-holotype).
The same characters cited for R. granuligera distinguish R. marysvillensis from
the treated species. Both species are closely related but they can be mainly
separated by the different ascospore size and shape being for R. marysvillensis
larger [(16-)18-22 x (8-)10-11(-13) um] and more broadly ellipsoid. Their
distribution it is also markedly different (see also AR. granuligera and R.
pruinella).
Distribution: Western North America (SHEARD in lit.).
Additional specimens examined: U.S.A.: California: Amador Co., on Alnus
rhombifolia, riparian woodland along Sutter Creek between Sutter Creek and
Volcano, ca. 2000 ft. alt, 15.3.1975, W.A. WEBER (SHEARD, aS WEBER: Lich.
exs. 474). San Francisco Co., San Francisco, Lands End, NE of Santa Cruz
Peninsula, on Alnus rubra, 25.6.1974, |.M. BRoDO 20481 & R.M. BROWN
(CANL).
Rinodina murrayii H. MAYRHOFER, Lichenologist 15(3): 273 (1983). - Type:
New Zealand: South Island, Otago, Lee Stream Valley south west of Dunedin,
2 km north of Lee Stream School, ca. 490 m, 23.9.1981, H. MAYRHOFER 2199
(GZU-holotype; CHR, OTA-isotypes).
The saxicolous habitat together with the Physcia-type ascospores make this
species distinct from its relatives (see R. santorinensis).
Distribution: New Zealand and Australia (MAYRHOFER 1983, 1984).
The specimens examined are listed in MAYRHOFER (1983).
Rinodina thujae (H. MAGN.) SHEARD, Bryologist 90: 164 (1987); A.
marysvillensis var. thujae H. MAGN., Bot. Not.: 192 (1953).- Type: U.S.A::
Wisconsin, Villas Co. Eagle River on Thuja occidentalis, 1946, J.W. THOMSON
2122 (UPS-holotype).
58
Only the larger areolae which do not dissolve into blastidia or soredia
distinguish this species from the closely related R. excrescens and R.
granulans (see comments made for these taxa).
Distribution: Great Lakes States and Canada (SHEARD in lit.).
Additional specimens examined: U.S.A.: Minnesota, St. Louis Co., Voyageurs
National Park, 11.6.1978, C.M. WETMORE 32864 (MIN).
We are indebted to the directors and keepers of the following herbaria: BM,
CANL, GZU, LISU, MIN, S, STU, TUR, W and WU; to Dr. O. BREuss (Vienna),
Dr. J. ETAYO (Navarra) and Dr. J. HAFELLNER (Graz) who kindly lent us their
private collections; to Mag. M. MATZER (Graz) for selecting specimens in BM; to
Prof. Dr. J.W. SHEARD (Saskatoon) for his valuable information and comments
on some specimens as well as for the loan of some specimens and correcting
the English text and to Prof. Dr. J. POELT (Graz) for critical revision of the
manuscript. The second author (H. MAYRHOFER) was supported by the Fonds
zur Forderung der wissenschaftlichen Forschung (Projekt P8500-BIO).
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Volume L, pp. 61-72 January-March 1994
SYSTEMATIC AND BIOLOGICAL STUDIESIN THE BALANSIEAE
AND RELATED ANAMORPHS. VIII.
THEEPHELIS ANAMORPH OF BALANSIA EPICHLOE.
RYAN A. PHELPS and GARETH MORGAN-JONES
Department of Plant Pathology, College of Agriculture and Alabama
Agricultural Experiment Station, Auburn University, Auburn, Alabama 36849
ABSTRACT
The development and morphology of the Ephelis state of Balansia
epichloe (Weese) Diehl on the adaxial surface of leaves of Eragrostis
capillaris (L.) Nees are described. Differences between the conidiomata of
this and those of some other Balansia anamorphs are noted. A formal taxonomic
description is presented.
INTRODUCTION
As discussed previously (Phelps et al., 1993b), Ephelis conidiomata of
Balansia species vary considerably in morphology, ranging from effuse and ill-
defined to distinctly hysteriform or apothecoid. Such structures have been
variously referred to as being pycnidial, sporodochium-like, patellate, or
cupulate (Diehl, 1950; Ullasa, 1969). Whatever their form, the conidiomata are
produced on primary stromata of diverse thicknesses, the latter
prosenchymatous to pseudoparenchymatous in texture and developing
superficially on host plant surfaces. Primary stromata of the various species
differ morphologically, depending upon the plant part on which they are
produced. Generally, when occurring on culms as in Balansia aristidae (Atk.)
Diehl, a pathogen of Aristida spp. (Phelps et a/., 1993a), and especially on
leaves as in Balansia epichloe, anamorph stromata are flat and effuse. When
associated with inflorescences, as in Balansia cyperi Edg. on Cyperus spp. and
Atkinsonella hy poxylon (Peck) Diehl on Danthonia spicata (L) P. Beauv. ex
Roem. & Schult., Ephelis conidiomata are often borne on somewhat pulvinate or
cushion-like stromata and are cupulate or patellate in form (Diehl, 1950;
Rykard et al., 1984; Leuchtmann and Clay, 1988).
Alabama Agricultural Experiment Station Journal Series Number 18-933559
62
Primary stromata formed on leaves, such as those of Balansia epichloe
and B. henningsiana (Moell.) Diehl, originate from hyphae egressing between
epidermal cells or through stomata (Phelps et a/., 1993c), and are usually thin,
diffuse, often extending and spreading some distance over the surface. These
two species, each of which occurs on a number of grass hosts, have often been
confused, but differ in that they form ascostromata that are epiphyllous and
hypophyllous, respectively. Balansia henningsiana occurs commonly in the
southern United States on species of Andropogon L., and Panicum L. (Diehl,
1950). Balansia epichloe has a similar geographical distribution and a host
range including species of Eragrostis Wolf, Panicum, and Sporobolus R. Br.
(Diehl, 1950).
In B. epichloe, hyphal egress through the adaxial leaf surface occurs
predominantly between epidermal cells and bulliform cells of the longitudinal
grooves (Phelps, et al., 1993c). At first, external hyphal proliferation occurs
primarily along the length of the longitudinal grooves proximal to the sites of
egress. This gives rise to a number of dense, linear mycelial mats in the
grooves, interconnected by sparse hyphae overlying the costal regions.
Eventually the primary stroma extends more or less evenly over the affected
portion of the leaf surface. The Ephelis anamorph is formed from this
stroma.
Rykard et al. (1984) briefly described the conidiomata of Atkinsonella
hypoxylon and several Balansia species, including B. epichloe and B.
henningsiana. Species were found to differ in a number of characteristics,
including extent of fertile conidiophore palisade development, thickness and
persistence of overlying stromal tissue, presence or absence of distinguishable
peripheral excipular tissue and spatial relationship of primary and secondary
perithecial stromata. In B. henningsiana, collected in Georgia on Andropogon
virginicus L., conidiomata were found to develop from long, narrow, thin, white
primary stromata formed on either or both sides of the midrib on the adaxial
leaf surface (Rykard et al., 1984). Palisades of conidiophores were reported
to develop in well-defined, irregular, hysteriform "structures" scattered on the
stroma surface before the leaf blade expanded. Perithecial stromata were
formed on the abaxial surface of the leaf blade opposite the conidial stromata.
Diehl (1950), in describing the ephemeral anamorph of B. henningsiana, made no
mention of occurrence on the upper leaf surface but noted that the stromata
were found on the abaxial surfaces of unrolling leaves. He referred to the
stroma as a "hypothallus". The conidiomata of B. henningsiana were described
as being linear to hysteriform, with thin, evanescent exciples.
PLATE 1. Leaf segments of Eragrostis capillaris showing dark Ephelis
conidiomata of Balansia epichloe on white, adaxially-borne primary stromata [X
5 to X 20]. A, young, more or less discrete, buff-colored fertile areas
(indicated by arrowheads); B, older, coalescent conidiomata assuming a linear
morphology; C-F, fully developed conidiomata; C, virtually covering entire
stroma; D&E, oblong to hysteriform, separated by sterile stromal mycelium,
with developing conidiomata (indicated by arrowheads) in close proximity; F,
variously confluent, giving an irregular to long-linear aspect.
63
64
Conidiomata of B. epichloe occurring on Chasmanthium laxum (L.)
Yates, were reported by Rykard et a/., (1984) to be in the form of irregular,
hysteriform, coalescing locules bordered laterally by ridges of sterile tissue.
When viewed in cross-section the fertile areas were described as having the
appearance of saucer-shaped depressions in the stroma. Diehl (1950) noted that
conidiomata of B. epichloe cover the so-called hypothallus as a whitish film that
splits irregularly in elongate, subhysteriform, coalescing pycnidia. These were
described as being partly separated by "the dissepiments of the surface layer as
peridia".
As already noted (Phelps et a/., 1993b), Diehl (1950) madedifferences in
conidiomatal and ascostromatal morphology the basis for recognition of two
subgenera of Balansia, namely Eubalansia and Dothichloe. The first was
characterized by possession of patellate or cupulate conidiomata with a well-
defined margin or excipulum, the second by effuse or hysteriform, more
irregular and less clearly defined, fertile areas. The utility of, and need for,
these subgeneric taxa have recently been questioned (Morgan-Jones et al., 1992).
Balansia is not a very large genus and its subdivision on this basis seems of
doubtful value.
The anamorph genus Ephelis Fr., based on E. mexicana Fr., contains
very few species although Ephelis states are known to be produced by most
Balansia species currently recognized, including B. aristidae, B. strangulans and
the others mentioned above. The affinity of the type species itself is uncertain
(Diehl, 1950; Phelps et a/., 1993b). It could be the anamorph of either Balansia
obtecta Diehl or B. claviceps Speg., probably the former. The binomial Ephelis
mexicana must be considered a nomen dubium because it cannot be properly
typified. Ephelis borealis Ell. & Everh. is the anamorph of Atkinsonella
hypoxylon (Peck.) Atk. and E. trinitensis Cke & Massee the anamorph of B.
claviceps. The status of Ephelis japonica P. Henn. and E. oryzae Syd. is also
uncertain since there seems little difference between the two (Phelps et al.,
1993b) and a teleomorph connection is not known for either.
Our collection and recent studies of several members of the Balansieae
in the southeastern United States (Morgan-Jones and White, 1992; Morgan-Jones
et al., 1993; Phelps et al/., 1993a, 1993b, 1993c) have been aimed at better
understanding host-pathogen relationships and documenting in greater detail the
characteristics of the respective fungi. The relationship between B. epichloe
and Eragrostis capillaris (L.) Nees and E. elliottii was reported upon in part
VII of this series (Phelps et al., 1993c). In the present paper, the development
and morphology of the Ephelis anamorph of B. epichloe on leaves of E.
capillaris are described.
MATERIALS AND METHODS
Test plants used during these studies were collected and maintained as
described by Phelps et al. (1993c). Leaf segments bearing ephelidial stromata
at various stages of development were harvested and prepared for sectioning.
Fixation, dehydration, infiltration, sectioning, and staining procedures were
performed as described by Phelps et al. (1993a).
Ss
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= —— S) AO e ©
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epichloe.
idiomata of Balansia
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FIGURE 1. Section
66
RESULTS
Primary Stroma Development
As documented in a preceding paper of this series (Phelps et a/., 1993c),
endophytic hyphae of Balansia epichloe grow systemically in host leaves,
longitudinally for the most part, between cells of both vascular and ground
tissue. Prior to stroma initiation, mycelium is relatively scattered and sparse.
It is most commonly associated with vascular bundles, particularly those of the
midvein. Primary stroma development and Ephelis anamorph production may
occur on mature, fully expanded leaves or when young and concealed by
enveloping leaves. Localized build-up of endophytic hyphae at a particular
position along the leaf blade, usually toward the middle, lengthwise, immediately
precedes initial hyphal egress. Stroma development follows at the same site.
As the area of hyphal concentration spreads in both directions, concurrent
egress from the adaxial leaf surface and stroma initiation occurs, both at first
along longitudinal grooves (Plate 2, B-D). Hyphal build-up, egress, and stroma
formation begin sequentially at the same locus, and subsequently spread up and
down the leaf blade. Initially, most hyphae egress between cells of the
longitudinal grooves. Small, parallel bundles of hyphae then grow linearly
within these intercostal grooves (Plate 2, B-D), eventually filling and
overflowing them (Plate 2, F). The stroma appears striate in its youngest
stages due to concentration of the delicate hyphal network within longitudinal
grooves (Phelps et a/., 1993c). In its final dimensions, the primary stroma may
cover the width of the blade and extend appreciably in length (Plate 1, C).
Eventually the dense, white, cottony stroma thickens, covering the entire width
of the affected leaf surface area (Plate 1, A &D). When mature, the primary
stromata assume a cream to very pale buff coloration and become more or less
prosenchymatous in texture. Constituent hyphae tend to be more densely packed
in the vicinity of longitudinal grooves.
Conidiomata develop at or near the surface of the stroma as elliptic to
hysteriform to linear pustules (Plate 1, A-F). Conidiophores originate from
hyphae occurring a short distance below the stromal surface. At first, the
developing palisades of conidiophores are covered by thin wefts of hyphae which
are soon disrupted, exposing the fertile areas. Conidiophore palisade primordia
are initially scattered (Plate 1, A) but often become progressively longer and
coalesce to form long, linear entities (Plate 1, B) following the subtending
longitudinal leaf grooves where the stroma is thickest and there is a
PLATE 2. Portions of thin sections showing developing primary stromata of
Balansia epichloe on adaxial leaf surfaces of Eragrostis capillaris plants. A,
cross section showing young, loose stroma (s) developing above two leaf veins
and in the longitudinal groove (indicated by arrowhead) between them [X 400];
B-E, cross sections showing young, loose stromata (s) and hyphae concentrated
within longitudinal grooves (indicated by arrowheads); (hyphae egressing
between epidermal cells indicated by arrows in D & E) [X 1000]; F, cross
section showing part of a more mature primary stroma (s) filling a longitudinal
groove [X 400].
68
preponderance of underlying, egressing hyphae. The palisades of conidiophores
rest ona loose mat of hyphae (Plate 3, D-F) and are essentially acervuloid in
aspect although the fructification is entirely superficial. When mature, the
conidiomata have a concave, saucer-like form when viewed in cross section.
Very young conidiomata are barely distinguishable from the surrounding sterile
stromal areas but with the initiation of conidiogenesis become discernible as
pale beige to very pale chocolate-colored patches (Plate 1, A). As conidiation
progresses, and mounds of conidia accumulate, the conidiomata assume a dark,
grayish-brown coloration. Conidiomatal coalescence and profuse conidium
production sometimes result in most of the primary stroma becoming covered by
dark masses of conidia (Plate 1, C). Under favorable conditions the fertile
areas become distinctly pustular in appearance due to the superficial
accumulation of conidial masses (Plate 1, D-F).
Taxonomic Description
Ephelis anamorph of Balansia epichloe (Weese) Diehl, USDA Agric. Monogr.
4:40, 1950.
Mycelium immersed in host leaf tissue, composed of hyaline, septate,
smooth, branched, 1.5-2 pm wide hyphae, distributed predominantly in the
vicinity of vascular bundles and immediately surrounding mesophyll. Hyphae
egressing predominantly between epidermal cells (Plate 2, D & E) or through
stomata. Primary stroma effuse to diffuse, especially toward the margin,
superficial, thin, at first somewhat lanose (Plate 1, E), becoming felted,
loosely prosenchymatous when mature, denser where close to the leaf surface,
particularly in proximity to longitudinal grooves, up to 120 ym thick, up to 2.5
cm in length, 2-6 mm wide, white when young, cream to very pale buff when
mature. Conidiomata eustromatic, as localized, undifferentiated, fertile areas
originating closely beneath the stroma surface, ephemeral, erumpent from
below a narrow layer of loosely interwoven hyphae, initially scattered,
becoming gregarious or sometimes confluent, elliptic to oblong to hysteriform
or long-linear in outline (Plate 3, D& F), acervular, saucer-shaped (Plate 3,
D), bordered peripherally by a small cluster of closely appressed, vertically
oriented, sterile hyphal cells as a rudimentary exciple (Figure 1), separated and
bordered by sterile strips of the primary stromata, about 1 mm wide, of various
lengths up to 1 cm. Conidiophores (Figure 2) micronematous, hyaline, smooth,
septate, simple or branched, arranged in dense fascicles, formed from hyphae
lining the base of the conidiomata. Conidiogenous cells long-cylindrical,
discrete, terminal or lateral, attenuating slightly toward the distal end, straight
or slightly flexuous, holoblastic, proliferating sympodially, indeterminate, giving
rise to a sequence of a few conidia at the apex, 1-1.5 pm wide. Conidia hyaline,
aseptate, narrowly fusiform, guttulate, more or less straight or slightly curved,
thin-walled, smooth, obtuse at the apex, subtruncate at the base, (16)-18-(21) X
1 pm (Figure 2).
On a number of Poaceae genera, including Andropogon L., Calamagrostis
Adams., Chloris Sw., Ctenium Panz., Eragrostis Wolf, Gymnopogon P. Beauv.,
Panicum L., and Sporobolus R. Br.; North, Central, and South America.
70
Collections examined: on Eragrostis capillaris, Auburn, Lee County,
Alabama, August 1991, G. Morgan-Jones, AUA; on E. elliottii, Geneva County,
Alabama, July 1991, R.A Phelps, AUA.
DISCUSSION
Although Balansia epichloe and B. henningsiana overlap in host range,
both occurring on Andropogon scoparius Michx., Gymnopogon ambiguus
(Michx.) B.S.P. and Panicum agrostoides Spreng. in the United States (Diehl,
1950), and are of broadly similar morphology, they are considered distinct
species. Not surprisingly, their superficial resemblance has led in the past to
their being regarded as a single entity. Ina key to species of Balansia, Diehl
(1950) separated the two taxa on the basis of their fructifications being
produced on opposite leaf surfaces; on the adaxial side in the case of B.
epichloe, on the abaxial side in B. henningsiana. Hyphal egress can, however,
apparently occur in both species from either leaf surface. Phelps et al. (1993c)
reported outward passage of hyphae of B. epichloe between epidermal cells on
the abaxial leaf in Eragrostis capillaris. Such hyphae do not continue to grow
and give rise to primary stromata however. As already mentioned, Rykard et
al. (1984) found the anamorph state of B. henningsiana formed on the adaxial
surface of leaves of Andropogon virginicus. Ascostromata of the two species
invariably occur on the different surfaces, in B. epichloe by conversion of the
primary stroma and development of perithecial initials a short distance below
the conidiophore palisade. From the account by Diehl (1950), and bearing in
mind the observations made by Rykard et al. (1984), it seems that B.
henningsiana can form primary stromata bearing conidiomata on either leaf
surface but further investigation of this species on different hosts is required
for purposes of confirmation.
The conidiomata of B. epichloe differ from those of species of Balansia
having effuse stromata occurring on culms, such as B. aristidae (Atk.) Diehl
(Phelps et a/., 1993b), in being much less well-defined and differentiated. The
primary stroma of B. aristidae becomes melanized at or about the time of
conidiomatal initiation and assumes a _ pseudoparenchymatous _ texture.
Conidiomata originate as locules in the stroma, the palisade of conidiophores
becoming exposed by a longitudinal split in the overlying tissue.
PLATE 3. Portions of thin sections showing developing primary stromata of
Balansia epichloe on adaxial leaf surfaces of Eragrostis capillaris plants. A,
oblique section showing bulliform cells (b), hyphae packed in longitudinal
groove (indicated by arrowhead), and overlying stroma (s) [X 500]; B & C,
cross sections showing Ephelis palisades (indicated by arrows) beginning to
form on underlying primary stromata (s) [X 300]; D-F, different magnifications
of a slightly oblique section showing a primary stroma (s) bearing a fully
developed Ephelis conidioma (indicated by arrowheads) [D; X 150; E; X 200; F;
X 400]. Underlying plant tissue includes bulliform cells (b) associated with a
longitudinal groove, and vascular tissue (v) surrounded by dark bundle sheath
cells.
71
72
ACKNOWLEDGEMENTS
We thank Dr. Mary E. Palm, USDA, Beltsville, Maryland, for her
presubmission review of this paper. The research on which it is based was
supported by a grant from the National Science Foundation (BSR-8922157).
LITERATURE CITED
DIEHL, W.W. 1950. Balansia and Balansieae in America. Agric. Monograph No.
4, USDA, Washington, D.C. 82 pp.
LEUCHTMANN, A. and K. CLAY. 1988. Atkinsonella hypoxylon and Balansia
cyperi, epiphytic members of the Balansieae. Mycologia 80: 192-199.
MORGAN-JONES, G. and J.F. WHITE, JR. 1992. Systematic and biological
studies in the Balansieae and related anamorphs. II. Cultural
characteristics of Atkinsonella hypoxylon and Balansia_ epichloe.
M ycotaxon 44: 89-102.
MORGAN-JONES, G., R.A. PHELPS and J.F. WHITE, JR. 1992. Systematic
and biological studies in the Balansieae. I. Prologue. Mycotaxon 43:
401-415.
MORGAN-JONES, G., R.A. PHELPS and M.R. OWSLEY. 1993. Systematic
and biological studies in the Balansieae and related anamorphs. VI. The
teleomorph of Balansia aristidae. Mycotaxon49: 107-116.
PHELPS, R.A., G. MORGAN-JONES and M.R. OWSLEY. 1993a. Systematic
and biological studies in the Balansieae and related anamorphs. IV.
Host-pathogen relationship of Aristida purpurascens and Balansia
aristidae. Mycotaxon 48: 165-178.
PHELPS, R.A., G. MORGAN-JONES and M.R. OWSLEY. 1993b. Systematic
and biological studies in the Balansieae and related anamorphs. V. The
Ephelis anamorph of Balansia aristidae. Mycotaxon 49: 91-105.
PHELPS, R.A., G. MORGAN-JONES and M.R. OWSLEY. 1993c. Systematic
and biological studies in the Balansieae and related anamorphs. VII.
Host-pathogen relationship of Eragrostis capillaris and Balansia
epichloe. Mycotaxon 49: 117-127.
RYKARD, D.M., E.S. LUTTRELL and C.W. BACON. 1984. Conidiogenesis and
conidiomata in the Clavicipitoideae. Mycologia 76: 1095-1103.
ULLASA, B.A. 1969. Balansia claviceps in artificial culture. Mycologia 61:
572-579.
MY COTAXON
Volume L, pp. 73-80 January-March 1994
A new species of Ascotricha with
non-ostiolate ascomata
Shun-ichi UDAGAWAD, Shigeru UCHIYAMA2) and
Seigo KAMIYA2)
1) Nodai Research Institute, Tokyo University of Agriculture,
1-1-1, Sakuragaoka, Setagaya-ku, Tokyo 156, Japan;
2) New Drug Discovery Research Laboratories, Tsukuba Research
Institute, Banyu Pharmaceutical Co., Ltd.,
3, Ookubo, Tsukuba-shi, Ibaraki 300-33, Japan
Abstract
Ascotricha distans, a new species with a Dicyma
anamorph, is described and illustrated from soil in
Japan. It is unique because of the production of
non-ostiolate ascomata, few ascomatal hairs and
evanescent asci, adaptive characters for its terrestrial
habitat. The Dicyma anamorph is characterized by
short conidiophores without sterile ampulliform
processes, geniculate, rachis-like conidiogenous cells
and pale, subglobose, almost smooth conidia.
Key Words Ascotricha , Dicyma, cleistothecial ascomycete,
Xylariaceae, taxonomy, soil fungus
The genus Ascotricha Berk. now contains 9 accepted species,
all of which have anamorphs classified in the form-genus Dicyma
Boulanger (Hawksworth, 1971; Kulshreshtha et al., 1977; von
Arx, 1981). Most species of Ascotricha are probably cosmopolitan
in distribution, and are commonly isolated from cellulosic
substrates (e.g. paper, wood, fabrics, plant debris, seeds, dung,
etc.), particularly in tropical to subtropical regions (Hawksworth,
1971; Hanlin, 1990). The occurrence in soil suggests an active
role in decomposing processes of cellulosic materials such as litter.
74
During our exploratory survey of soil-borne ascomycetes as
producers of secondary metabolites useful to the pharmaceutical
industry, an interesting member of Ascotricha was isolated from
cultivated soil in a herbal plant garden, southern Kyusyu. The
fungus superficially resembles A. amphitricha (Corda) Hughes
(Hawksworth, 1971) in morphology of ascomatal hairs, but differs
from it in producing non-ostiolate ascomata with few hairs and
clavate asci, aS well as in its conidial characters. Recently,
Valldosera and Guarro (1988) erected the monotypic genus
Ascotrichella, which was considered as a transitional form of the
Xylariaceae between Ascotricha and Coniochaeta (Sacc.) Cooke.
Ascotrichella hawksworthii Valldosera et Guarro is characterized by
short hairy ostiolate ascomata, cylindrical asci with a non-amyloid
apical apparatus, discoid ascospores with a longitudinal germ slit
and a Humicola-like anamorph. Our fungus distinctly differs from
Ascotrichella hawksworthii in having long, distinct, smooth-walled
hairs and a Dicyma anamorph. Thus, it is described as a new
species of Ascotricha herein.
Ascotricha distans Udagawa, Uchiyama et Kamiya, sp.nov.
Figs:
Coloniae in agaro cellulosa restrictae, planae, tenues, ex
mycelio vegetativo submerso et hyphis aeriis sparsis formantes,
brunneo-griseae vel ferro-griseae; ascomata tarde producentia, nigra;
conidiogenesis dispersa; reversum fumosum.
Ascomata superficialia vel immersa, dispersa, non-ostiolata,
nigra, nitida, globosa vel subglobosa, 70-120 um _ diam,
paucipilosa; pili dimorphi: (a) longi, flagelliformes, atrobrunnei,
rigidi, simplices, 500-1000 x 2-2.5 um, basi 5-7.5 Um diam,
angustati, leves, septati, raro cum ramis ampulliformibus instructi,
(b) setiformes, saepe deminuentes, aseptati vel pauciseptati, abrupte
angustati, ad apicem interdum cellula ampulliformi hyalina
formantes; peridium olivaceo-brunneum, tenue, ex “textura
epidermoidea” compositum. Asci 8-spori, clavati, 30-42.5 x 7.5-10
uum, brevi-stipitati, evanescentes; paraphyses nullae. Ascosporae
biseriatae, primum hyalinae, postremo valde olivceo-brunneae,
ovoideae vel ellipsoideae, compressae a latere visis, 5-6.5 (-7.5) xX
4.5-5 x 3-3.5 tum, leves, fissura germinali paratae. Status
anamorphus: Dicyma sp.
Holotypus BF 39285, colonia exsiccata in cultura ex solo
sativo, Kaimon-cho, Ibusuki-gun, Kagoshima, in Japonia, 28. 1x.
1992, a S. Uchiyama et S. Kamiya isolata et ea collectione
fungorum, Musei et Instituti Historiae Naturalis Chiba (CBM)
conservata.
Etymology: from Latin, distans=distinct, referring to the
distinct characters of ascomata.
Fig. 1. Ascotricha distans, BF 39285.
A. Ascomata; B. Portions of long ascomatal hairs; C. Seta-like
short ascomatal hairs; D. Asci; E. Ascospores; F. Conidiogenous
cells; G. Conidia.
be.
76
Anamorphosis: Dicyma sp.
Mycelium ex hyphis hyalinis vel dilute brunneis, ramosis,
septatis, levibus, 1-2.5 um diam compositum. Conidiophora erecta,
ex mycelio basali vel hyphis aeriis oriunda; stipites 10-50 x 1.5-3
uum, leves, incrassati, septati, simplices vel apicaliter verticillate
ramosi, basi olivaceo-brunnei, apicem versus gradatim pallescentes
et angustati. Cellulae conidiogenae ex conidiophoro simplici vel
ramis terminales, subhyalinae vel dilute brunneae, basi
ampulliformes vel elongatae, 8-12 x 3-4 tum, apice sympodice
prolongatae, usque 20-50 um longae, 2.5 -3 tm diam, geniculatae.
Conidia ex denticulis terminalibus successivis singulatim producta,
hyalina vel dilute olivaceo-brunnea, subglobosa, 3.5-5 um diam,
vel obovoidea, 3-5.5 x 2.5-5 tm, levia vel parum asperata, ad
basim cicatrice inconspicua affixa.
Holotypus BF 39285, loc. cit.
Colonies on cellulose agar growing restrictedly, attaining a
diam of 10-12 mm in 14 days at 25°C, plane, thin, consisting of
a submerged vegetative mycelium and sparse aerial hyphae,
Brownish Grey (M. 6F2, after Kornerup and Wanscher, 1978) or
Iron Grey (Rayner, 1970), later producing black ascomata on the
substratum; conidiogenesis sparse; reverse Smoke Grey (R).
Colonies on potato-carrot agar (PCA) growing rather restrictedly,
attaining a diam of 17-19 mm in 14 days at 25°C, velvety, plane
or slightly sulcate, consisting of a thin basal felt, Dark Green (M.
27F3) or Olivaceous Black (R), producing abundant ascomata
covered by a dense overgrowth of conidia; reverse Greenish Grey
(M. 27F2) or Olivaceous Black (R).
Ascomata superficial or sometimes immersed, often intermixed
with profuse conidiophores, scattered, non-ostiolate, black, shining,
globose to subglobose, 70-120 tum diam, with a few, long,
straight or slightly flexed hairs on the upper part and several short
ones the lower part; ascomatal hairs of two types: (a) long, whip-
like, dark brown, rigid, unbranched, 500-1000 um long, basally
5-7.5 um wide, tapering and 2-2.5 tm wide throughout most of
the length, smooth and thick-walled, septate, slightly swollen at
the septum, rarely with short hyaline ampulliform branches, and
(b) seta-like, often diminished, aseptate or few septate, abruptly
tapering to a pointed tip or a hyaline ampulliform cell measuring
7.5-15 x 3-5 jm; ascomatal peridium olivaceous brown, thin, of
textura epidermoidea; outer layer consisting of dark, thick-walled,
irregular cells measuring 5-7.5(-12.5) 4m diam, and inner layer of
hyaline, thin-walled, angular cells. Asci 8-spored, clavate, 30-
42.5 x 7.5-10 um (p. sp. 17.5-25 tm long), short-stipitate up to
7.5-12.5 um _ long, thin-walled, without apical structures,
evanescent; paraphyses not observed. Ascospores biseriate, at first
1s 4
Fig. 2. Ascotricha distans, BF 39285.
A, B. Asci, x 1200; C, D. Ascospores (SEM), x 2245(C) and
x 2500(D); E, F. Conidiogenous cells and conidia, x 1200.
78
hyaline, becoming dark olivaceous brown at maturity, ovoid to
ellipsoidal, compressed when seen edgewise, 5-6.5(-7.5) x 4.5-5
x 3-3.5 um, rounded at both ends, smooth-walled, with a
longitudinal germ slit.
Mycelium consisting of hyaline to pale brown, branched,
septate, smooth-walled, 1-2.5 tm _ thick hyphae. Conidiophores
erect, arising from the basal mycelium or aerial hyphae; stipes
short, 10-50 x 1.5-3 tm, smooth and thick-walled, septate,
unbranched or apically branched a few times in_ verticillate
arrangements, olivaceous brown at the base, gradually paling and
tapering towards the apex. Conidiogenous cells arising terminally
from unbranched conidiophore or branches of the conidiophore,
subhyaline to pale olivaceous brown, consisting of a flask-shaped
to elongate basal part, 8-12 x 3-4 Um, and a well-developed,
denticulate rachis to 20-50 um long and 2.5-3 um _ wide,
geniculate, with crowded, distinct, about 1 tm wide conidium-
bearing denticles. Conidia acropleurogenous, produced sympodially
on the denticle, hyaline to pale olivaceous brown, subglobose, 3.5-
5 um diam, or obovoid, 3-5.5 x 2.5-5 um, the walls smooth to
faintly roughened, with an inconspicuous basal scar.
At 37°C, growth is nil.
Specimen examined: BF 39285 (holotype), in dried culture
isolated from cultivated soil, herbal plant garden, Kaimon-cho,
Ibusuki-gun, Kagoshima-pref., Japan, 28 Sept. 1992, col. S.
Uchiyama and S. Kamiya. The holotype has been deposited with
the Natural History Museum and Institute, Chiba, Japan (CBM).
The genus Ascotricha is found in the earlier literature under
the Chaetomiaceae (Ames, 1963). On the basis of the distinct neck
on the perithecia, the smooth-walled terminal hairs, the amyloid
plug in the asci, the prominent germ slit in the ascospores and the
anamorphs close to Geniculosporium-Nodulisporium, it is
distinguished from Chaetomium and now best included in the
Xylariaceae (Hawksworth and Wells, 1973; Khan and Cain, 1977;
von Arx, 1982; Eriksson and Hawksworth, 1991). Ascotricha
distans is the first species of the genus known to produce
cleistothecia, with few hairs and evanescent asci. As in the
Ceratostomataceae, Chaetomiaceae Lasiosphaeriaceae, Microascaceae
and Sordariaceae, particularly in species found in soil, such
morphological adaptations are well-recognized by the result of a
progressive evolution away from dispersal of ascospores by means
of air currents. Although there are only a few examples in the
Xylariaceae (Malloch and Rogerson, 1977), the convergence in the
ascoma characters of A. distans is considered to be an adaptation
to its terrestrial habitat analogous to those in other cleistothecial
ascomycetes.
72
The Dicyma anamorph of new species, is not entirely typical
of Dicyma states of Ascotricha species as_ described by
Hawksworth (1971) and von Arx (1982) on account of the
absence of sterile ampulliform cells on the conidiophores. Of the
known species of Dicyma (von Arx, 1982; Guarro and Calvo,
1983), D. pulvinata (Berk. et Curt.) von Arx (=Hansfordia
pulvinata (Berk. et Curt.) Hughes) is most close to our fungus.
Dicyma pulvinata differs from the fungus in having long, slender
conidiophores with a_ well-developed branching system,
conidiogenous cells with a very short rachis and somewhat larger
conidia (Hughes, 1951; Tubaki, 1958; Ellis, 1971; Matsushima,
1975; von Arx, 1981).
We are grateful to Dr. Cannon, CAB _ International
Mycological Institute, for his critical reading of the manuscript.
Literature cited
Ames, L.M. 1961 [1963]. A monograph of the Chaetomiaceae.
Us SwvAnny: Res. “Dev; Ser. 2. 125p.
Arx, J.A. von. 1981. The genera of fungi sporulating in pure
culture, 3rd ed. J. Cramer, Vaduz. 424p.
Arx, J.A. von. 1982. The genus Dicyma, its synonyms and
related fungi. Proc. Kon. Ned. Akad. v. Wet., Amsterdam,
SeriC. $5: 21-28
Ellis, M.B. 1971. Dematiaceous Hyphomycetes. Commonwealth
Mycological Institute, Kew. 608p.
Eriksson, O.E. and Hawksworth, D.L. 1991. Outline of the
Ascomycetes 1990. Syst. Ascomycet., 9: 39-271.
Guarro, J. and Calvo, M.A. 1983. Dicyma funiculosa sp.nov.
from Spain. Nova Hedwigia, 37: 641-649.
Hanlin, R.T. 1990. Illustrated genera of Ascomycetes. APS Press,
St. Paul, Minnesota. 263p.
Hawksworth, D.L. 1971. A _ revision of the genus Ascotricha
Berk. Commonwealth Mycological Institute Mycol. Pap., 126:
1-28.
Hawksworth, D.L. and Wells, H. 1973. Ormamentation on the
terminal hairs in Chaetomium Kunze ex Fr. and some allied
genera. Commonwealth Mycological Institute Mycol. Pap.,
134: 1-24.
Hughes, S.J. 1951. Studies on micro-fungi. IX. Calcarisporium,
Verticicladium and Hansfordia (Gen. Nov.). Commonwealth
Mycological Institute Mycol. Pap., 43: 1-25.
Khan, R.S. and Cain, R.F. 1977. The occurrence of amyloid
plugs in the asci of Ascotricha erinacea. Mycotaxon, 5: 409-
414.
80
Kornerup, A. and Wanscher, J.H. 1978. Methuen handbook of
colours, 3rd ed. Eyre Methuen, London. 252p.
Kulshreshtha, D.D., Raychaudhuri, S.P. and Khan, A.M. 1977.
Studies on some soil fungi associated with maize (Zea mays
L.) 1. Three new Ascomycetes. Acta Bot. Indica, 5: 16-19.
Malloch, D. and Rogerson, C.T. 1977. Pulveria, a new genus of
Xylariaceae (Ascomycetes). Can. J. Bot., 55: 1505-1509.
Matsushima, T. 1975. Icones microfungorum a Matsushima
lectorum. Kobe, Japan. 209p. + 415 pl.
Rayner, R.W. 1970. A mycological colour chart. Commonwealth
Mycological Institute, Kew.
Tubaki, K. 1958. Studies on the Japanese Hyphomycetes. V.
Leaf & stem group with a discussion of the classification of
Hyphomycetes and their perfect stages. J. Hattori Bot. Lab.,
20: 142-244.
Valldosera, M. and Guarro, J. 1988. Some _ coprophilous
Ascomycetes from Chile. Trans. Br. mycol. Soc., 90: 601-
605.
MY COTAXON
Volume L, pp. 81-88 January-March 1994
TALAROMYCES SPECTABILIS, A NEW SPECIES OF
FOOD-BORNE ASCOMYCETES
Shun—ichi UDAGAWA' and Shoji SUZUKI“
1 Nodai Research Institute, Tokyo University
of Agriculture, 1-1-1 Sakuragaoka, Setagaya-
ku, Tokyo 156, Japan
2 Japan Food Research Laboratories, 52-1
Motoyoyogi-cho, Shibuya-ku, Tokyo 151, Japan
ABSTRACT
A new species of Talaromyces with a Pae-
cilomyces anamorph, T. spectabilis Udagawa et
S. Suzuki, isolated from heat processed fruit
beverage in Japan as a contaminant and from
road-side soil in Nepal, is described and il-
lustrated. The fungus is characterized by
fast growing, olivaceous brown colonies,white
ascomata, large, almost smooth-walled, ellip-
soidal ascospores, repeatedly branched coni-
diophores and cylindrical to ovoidal conidia.
A variety of heat resistant fungi are often found
in spoilage of fruit juices and other heat processed
beverages. The contamination of these products is gener-
ally caused by members of soil-borne ascomycetes such
as Byssochlamys, Eupenicillium, Hamigera, Neosartorya and
Talaromyces (Samson, 1989; Samson et al., 1992).
During a survey of mold spoilage of commercial
beverages in our laboratory, a number of heat resistant
fungi were isolated. Among these is a cleistothecial
ascomycete belonging to Talaromyces C, R. Benjamin (Stolk
and Samson, 1972). Based on the presence of a Paecilomyces
anamorph and its thermotolerant growth, it is classified
in the section Emersonii (Stolk and Samson, 1972) of the
genus but is clearly distinct from the two related spe-
cies described hitherto. A description of the fungus is
provided herein and a new name is proposed for it.
82
Talaromyces spectabilis Udagawa et S. Suzuki, sp.nov.
PLS cts
Coloniae in agaro '"Czapek-yeast extract (CYA)"
effusae, velutinae vel funiculosae, planae sed in centro
rugosae et sulcatae, ex mycelio basali coacto tenuiter
constantes, olivaceo-brunneae-_—ive | viridi-olivaceae;
ascomata nulla vel limitata; conidiogenesis profusa;
reversum griseo-flavum vel olivaceo-bubalinum. Coloniae
in agaro maltoso (MEA) effusae, velutinae vel funiculosae,
planae, ex mycelio basali coacto tenuiter constantes,
olivaceo-brunneae vel melleae vel viridi-olivaceae;
ascomata tarde producentia sed abundantia, alba; conidio-
genesis conferta; reversum incoloratum vel brunneo-auran-
tiacum vel melleum. Coloniae in agaro farinae avenaceae
effusae, velutinae vel plus minusve funiculosae, planae,
ex mycelio basali coacto tenuiter constantes, olivaceo-
brunneae vel viridi-olivaceae; ascomata tarde producentia,
alba; conidiogenesis profusa; reversum dilute flavum
vel primulinum.
Ascomata discreta vel interdum confluentia, non-
ostiolata, tarde maturescentia, alba, globosa vel subglo-
bosa, 100-350(-480) um diam, mollia, mycelio laxo stricto
vel sinuoso stramineo incrustato obtecta; paries ex
hyphis hyalinis vel dilute flavis ramosis septatis varie
asperatis intricatis compositus. Asci. singulariter
formantes, (4-)8-spori, subglobosi vel late ovoidei
vel pyriformes, 10-14 x 10-12 um, brevistipitati, evanes-—
centes. Ascosporae postremo stramineae, ellipsoideae,
5.5-7(-9) x 4-5.5 um, fere leves et incrassatae. Status
anamorphus: Paecilomyces spectabilis.
Ubiquinona principalia: Q-9 et Q-10.
Holotypus SUM 3030, colonia exiccata in cultura
ex potione pomo, Tokyo, in Japonia, xi.1992, a S. Suzuki
isolata et ea collectione fungorum, Musei et Instituti
Historiae Naturalis Chiba (CBM) conservata.
Etymology: from Latin, spectabilis=remarkable, refer-
ring to the wide-spread growth on common media.
Anamorphosis: Paecilomyces spectabilis Udagawa et S. Suzu-
ki, anam.nov.
Conidiophora ex mycelio basali vel hyphis aeriis
oriunda; stipites hyalini, 25-240 x 4-6(-8) um, leves sed
inferne manifesto asperati, septati. Penicilli complexi,
verticillati vel irregulatim ramosi, saepe divergentes,
interdum biverticillati vel monoverticillati. Rami
7-35 x 2.5-5(-7) um. Metulae 2-4 verticillatae, 8-10 x
2.5-3(-6) um. Phialides 2-8 verticillatae, 10-30 x 1.5-4
um. Conidia hyalina vel dilute flavo-brunnea, levia, pri-
Wt ?
=
Vy
U YF 7 ies ()
\ Gr
Figure 1. Talaromyces spectabilis, SUM 3030.
A. Asci. 8. Ascospores. . Ascomatal initials.
D. Conidiophor idiogenous cells. E. Conidia.
F, Chlamydospo
UDAGAWA & SUZUKI
84
mum cylindracea, 4-7 x 1.5-2(-3) um, utrinque saepe trun-
cata, postremo ovoidea vel ellipsoidea, 4-10 x 2-/ um, in
catenis laxe columnaribus vel implicatis connexa. Status
teleomorphus: Talaromyces spectabilis.
Holotypus SUM 3030, loc. cit.
Colonies on Czapek agar growing rapidly, attaining a
diameter of 38-52 mm within 7 days at 25°C, velvety
to more or less funiculose, plane, consisting of a thin
basal felt, Olive Brown (M. 4D6, after Kornerup and
Wanscher, 1978) or Honey to Citrine (Rayner, 1970);
ascomata usually absent; conidiogenesis profuse; odor
musty; reverse Greyish Yellow (M. 4C3) or Olivaceous
Buff (R). Colonies on CYA growing rapidly, attaining
a diameter of 65-75 mm within 7 days at 25°C, velvety
to funiculose, plane but centrally wrinkled and sulcate,
consisting of a thin basal felt, Olive Brown (M. 4E6)
or Greenish Olivaceous (R); margins broad and submerged;
ascomata absent or limited; conidiogenesis profuse;
exudate limited, clear; odor strongly musty; reverse
Greyish Yellow (M. 4C4) or Olivaceous Buff (R). Colonies
on MEA spreading broadly, attaining a diameter of /4-
85 mm or more within 7 days at 25°C, velvety to funicu-
lose, plane, consisting of a very thin basal felt, Olive
Brown (M. 4D5-E4) or Honey to Greenish Olivaceous (R),
with surface becoming dense-textured due to conidial
growth; margins broad and thin; ascomata slowly develop-
ing in an uneven layer beneath the conidial growth,
numerous in number, white; odor musty; reverse uncolored
to Brownish Orange (M. 5C4) or Honey (R). Colonies
on oatmeal agar spreading broadly, attaining a diameter
of 68 mm or more within 7 days at 25°C, velvety to more
or less funiculose, plane, consisting of a thin basal
felt, Olive Brown (M. 4E4) or Greenish Olivaceous (R);
ascomata slowly developing on the felt, white; conidio-
genesis profuse; reverse Pale Yellow (M. 1A3) or Primrose
(R). Colonies on cornmeal agar growing rapidly, attain-
ing a diameter of 60-62 mm within 7 days at 25°C, plane,
thin, vegetative mycelium submerged, with surface bearing
scattered conidia, Olive Brown (M. 4D5) or Honey (R);
ascomata limited; reverse uncolored.
Ascomata discrete or sometimes confluent, non-ostio-
late, maturing slowly within 30 days, white, globose
to subglobose, 100-350(-480) um in diam, soft, loosely
covered by straight or sinuous, straw-colored, encrusted
hyphae; ascomatal wall rather rudimentary, consisting
of a loose network of hyaline to light yellow, branched,
septate, often constricted at the septum, variously
(INAGAWA & SIIZIIKT y)
Figure 2. Talaromyces spectabilis, SUM 3030.
A. Asci. B. Ascospores. C, D. Conidiophores and
conidiogenous cells. E. Conidiogenous Geblise inh:
Conidia. Scale bars for A, B, E, F=10 um and for
Gacv=220'"uim,
85
86
roughened, interwoven hyphae measuring 2-4 um in diam.
Ascomatal initials arising as coiled or contorted side
branches of aerial hyphae, soon becoming surrounded
by hyphae arising from the neighboring cells. Asci
borne singly from fertile hyphae, (4-)8-spored, subglo-
bose to broadly ovoidal or pyriform, 10-14 x 10-12 um,
short-stipitate, evanescent. Ascospores' straw-colored
at maturity, ellipsoidal, 5.5-7(-9) x 4-5.5 um, more or
less roughened when young, almost smooth and thick-walled,
without ridges.
Conidiophores arising from the basal mycelium or
aerial hyphae; stipes hyaline, 25-240 x 4-6(-8) um, smooth-
walled but distinctly roughened in lower part, septate,
sometimes swollen at the septum. Penicilli (conidiogen-
ous cells) complex, consisting of a dense whorl of
verticillately or irregularly arranged, often divaricate
branches, sometimes biverticillate or monoverticil late.
Rami variable and often arranged at different levels,
divergent, sometimes recurved, 7-35 x 2.5-5(-7) um. Metu-
lae not readily distinguishable from the uppermost rami,
2-4 in the verticil, 8-10 x 2.5-3(-6) um. Phialides 2-8
in the verticil, 10-30 x 1.5-4 um, consisting of a cylind-
rical basal portion, tapering abruptly to a long distinct
neck, smooth-walled. Conidia hyaline to pale yellow-
brown, smooth-walled, variable in shape, cylindrical,
4-7 x 1.5-2(-3) um, with both ends truncated, becoming
ovoidal to ellipsoidal, 4-10 x 2-/ um, borne in loosely
columnar or disordered, long chains up to 200-300 um long
or more. Chlamydospores present, brown, globose or pyri-
form, 5.5-10 um in diam, smooth and thick-walled, either
terminal or intercalary. Racquet mycelium. present,
swollen near the septum up to 10-12 um in diam. Mycelium
consisting of hyaline or brown, branched, often thick-
walled, smooth or roughened, septate, 1.5-/.5 um wide
hyphae.
Main ubiquinones: Q-9 + Q-10.
At- 37°C. growths: slightly faster.
Specimens examined: SUM 3030 (holotype), in dried
culture isolated from heat processed fruit beverage
as a contaminant, Tokyo, Japan, November 1992, col.
S. Suzuki; and SUM 3031, in dried culture isolated from
road-side soil, Ring Road, Kathmandu, Nepal, 30 September
1986, col. S. Udagawa. The holotype has been deposited
with the Natural History Museum and Institute, Chiba,
Japan (CBM).
Talaromyces spectabilis is distinctive in several
respects. There is some superficial similarity between
87
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88
two described species of the genus, viz. T. byssochlamyd-
oides Stolk et Samson and T. leycettanus Evans et Stolk
in their thermo-resistant property and in the associated
Paecilomyces anamorphs (Evans and Stolk, 1971; Stolk
and Samson, 1972; Samson, 1974). From both species, Ty
spectabilis differs in having much larger ascospores
as well as in other several characters shown in Table
1. The Paecilomyces anamorph of T. spectabilis can be dis-
tinguished from Paecilomyces variotii Bainier (Samson, 19
74) in having larger conidium-bearing structures and
longer cylindrical conidia. The ubiquinone system of P.
variotii was determined as only Q-9 or Q-10 by Kuraishi
et al. (1985). This also supports a specific separation
of P. variotii from the anamorph of the new species.
Acknowledgement: We thank Prof. R.T. Hanlin of the Univ-
ercity of Georgia for reviewing this paper.
Literature cited
Evans, H.C. and Stolk, A.C. 1971. Talaromyces leycettanus
sp.nov. lpanss core imycol.) (Soceg, 565. 45-49.
Kornerup, A. and Wanscher, J.H. 1978. "Methuen handbook
of colour," 3rd ed. Eyre Methuen, London. 252p.
Kuraishi, H., Katayama-Fujimura, Y., Sugiyama, J., and
Yokoyama, T. 1985. Ubiquinone systems in fungi. I.
Distribution of ubiquinones in the major families
of ascomycetes, basidiomycetes, and deuteromycetes,
and their taxonomic implications. Trans. Mycol.
Soc. Japan, 26%) 383-395:
Kuraishi, H., Aoki, M., Itoh, M., Katayama, Y., Sugiyama,
J. and wPT bovis 99 lee wDistribution ofeub duds
nones in Penicillium and related genera.Mycol. Res.,
95: 705-711.
Rayner, R.W. 1970. "A mycological colour chart." Common-
wealth Mycological Institute, Kew.
Samson, R.A. 1974. '"Paecilomyces and some allied Hypho-
mycetes.'' Studies in Mycology No. 6. Centraalbureau
voor Schimmelcultures, Baarn. 119p.
Samson, R.A. 1989. Filamentous fungi in food and feed.
J. “Apple Bacteriol. Syme) Suipp te) 67 82/7 S-655.
Samson; R. AvyeHocking, “A.D, «Pitt, d7 10s andi Kings AsU:
(ed.). 1992. "Modern methods in food mycology."
Elsevier, Amsterdam. 388p.
Stolk, A.C. and Samson, R.A. 1972. "Studies on Talaromy-
ces and related genera II. The genus Talaromyces."
Studies in Mycology No. 2. Centraalbureau voor
Schimmelcultures, Baarn. 56p.
MY COTAXON
Volume L, pp. 89-92 January-March 1994
NOTES ON THE GENUS MONOICOMYCES
(LABOULBENIALES, ASCOMYCOTINA):
ON THE SPECIES DESCRIBED BY SPEGAZZINI
SERGIO SANTAMARIA
Unitat de Botanica. Departament de Biologia Animal, Biologia Vegetal i Ecologia.
Facultat de Ciéncies. Universitat Autonoma de Barcelona.
08193-Bellaterra (Barcelona). Spain.
ABSTRACT
In order to clear up the taxonomic status of Monoicomyces ternatus, Monoicomyces
unilateralis and Monoicomyces venetus their types have been studied. M. ternatus and
M. unilateralis are here proposed as new synonyms for Monoicomyces homalotae. M.
venetus is retained as a separate species.
Spegazzini described six species of Monoicomyces: M. infuscatus Speg. on
Gyrohypnus bonariensis Gemm. & Har. (=G. gracilis, as Xantholinus gracilis Boh.)
from Argentina (Spegazzini, 1912); M. affinis Speg. on an undetermined staphylinid
from Italy; M. ternatus Speg. on a mycophilous staphylinid from Italy; M. unilateralis
on an undetermined staphylinid from Italy; M. venetus Speg. on an undetermined
staphylinid from Italy (Spegazzini, 1915); and M. ocaleae Speg. on Ocalea funebris
Lynch from Argentina (Spegazzini, 1917).
Monoicomyces affinis was included as synonymous with Monoicomyces invisibilis
Thaxt. by Thaxter (1931) and this disposition has been subsequently accepted by all the
authors who mention M. invisibilis. Monoicomyces ocaleae is a synonym of
Monoicomyces caloderae Thaxt. according to Thaxter (1931). I have studied the type
of M. ocaleae, and my own observations are in full agreement with this opinion.
Monoicomyces infuscatus is a well defined species strictly parasitic on Xantholinini
staphylinids. However, the chief purpose of this note is to comment on the other three
species described by Spegazzini: M. ternatus, M. unilateralis and M. venetus.
These three species were described in the same work (Spegazzini, 1915) and no
further records have been added. To clear up the taxonomic status of these species I
requested the Institute of Botany "C. Spegazzini" in La Plata, Argentina, to lend me
their types for examination.
Monoicomyces ternatus Spegazzini, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 67 (1915)
The slide examined is labelled as follows: "STAFILINO MICOFILO / CONEGLIANO /
90
XI-1914 / LPS 45061 / SINTIPO".
This slide only contains one mature specimen (fig. 1). According to the description
of Spegazzini the most reliable characteristics of this species were the long secondary
appendages with clavate cells. As in M. unilateralis and M. venetus, their descriptions
are too brief and the drawings not very clear. My observations suggest that M. ternatus
is a form of M. homalotae, accepted as a variable and rather polymorphic species.
The primary appendage appears to be broken above its second cell. The two basal
cells of the primary appendage are deeply brown-pigmented and are separated by a pale
septum, as is typical in M. homalotae. The antheridial and perithecial features are not
distinct from those included in the wide range of characters of M. homalotae.
Therefore, because of the absence of any outstanding characteristic, I concluded that
M. ternatus should be a synonym of M. homalotae.
Monoicomyces unilateralis Spegazzini, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 68 (1915)
The slide examined is labelled as follows: "ATHETA ? ... MICOFILA / CONEGLIANO
/ 1X-1914 / LPS 45087 / Tipo".
This slide contains 13 immature or nearly mature specimens (figs. 2-4). Like M.
ternatus, this species was considered to be related to Monoicomyces britannicus Thaxt.
by Spegazzini. The most important characteristics were those of the primary appendage
and the absence of secondary appendages on the antheridium. Undoubtedly, the
specimens seen belong in M. homalotae, a species often confused with M. britannicus
(Santamaria, 1989). Specimens of M. unilateralis have characteristics that are very
typical in M. homalotae: there is a heart-shaped cell I and the base of the primary
appendage is deeply brown-pigmented with a characteristic pale ring between the two
basal cells. The secondary appendages may be present or be inconspicuous (unicellular)
in M. homalotae, this character being very variable. The supposed “unilaterality" of the
thallus in this species is not consistent, and normal, bilaterally symmetric thalli are
abundant among the specimens included on the type slide (fig. 4). Therefore, I conclude
that M. unilateralis should be regarded as a synonym of M. homalotae.
Monoicomyces venetus Spegazzini, Anales Mus. Nac. Hist. Nat. Buenos Aires 27: 69 (1915)
The slide examined is labelled as follows: "IN CAPITE STAFILIN ... / CONEGLIANO
/ VII-1913 / LPS 45062 / Tipo".
This slide contains only one mature specimen (fig. 5). This species is very different
from the other two herein studied. Monoicomyces venetus seems to be a well
distinguished species: primary appendage unbranched, short, and amber-coloured as in
the remainder of the thallus; perithecium with the outer walls arranged in spiral rows;
base of the perithecial stalk cell dark and constricted. Therefore, this species may be
Fig. 1. Monoicomyces ternatus (LPS:45061). Sintype. Figs. 2-4. Monoicomyces
unilateralis (LPS-45087). Type specimens. Fig. 5. Monoicomyces venetus (LPS:45062).
Type specimen. (Scale bars = 50 um. Scale bar in fig. 2 is the same for the figs. 3,
4 and S.)
91
92
retained as a separate species.
The author wishes to express his gratitude to Angélica Arambarri (director of “Instituto de Botanica
C. Spegazzini") for the loan of Spegazzini types, and to I. Tavares for critical reading of the manuscript.
REFERENCES
Santamaria, S. 1989. El orden Laboulbeniales (Fungi, Ascomycotina) en la Peninsula Ibérica e Islas
Baleares. Edicions especials de la Societat Catalana de Micologia. Vol. 3. 396 Pp. Esplugues de
Llobregat.
Spegazzini, C. 1912. Contribucién al estudio de las Laboulbeniomicetas argentinas. Anales Mus. Nac.
Hist. Nat. Buenos Aires 23: 167-244.
Spegazzini, C. 1915. Segunda contribucién al conocimiento de las Laboulbeniales italianas. Anales Mus.
Nac. Hist. Nat. Buenos Aires 27: 37-74.
Spegazzini, C. 1917. Revisién de las Laboulbeniales argentinas. Anales Mus. Nac. Hist. Nat. Buenos
Aires 29: 445-688.
Thaxter, R. 1931. Contribution towards a monograph of the Laboulbeniaceae. Part V. Mem. Amer.
Acad. Arts Sci. 16: 1-435.
MYCOTAXON
Volume L, pp. 93-98 January-March 1994
OBSERVATIONS ON PSEUDOTRACYLLA SPECIES
L. M. CARRIS
Department of Plant Pathology
Washington State University
Pullman, Washington 99164-6430
ABSTRACT
Pycnothyrial formation and sporulation in Pseudotracylla dentata
is described from a collection on dead leaves of Vaccinium macrocarpon
(cranberry) from a cultivated bog in Massachusetts. This is the first report
of P. dentata since its description in 1976 on Eucalyptus leaves in Brazil.
The geographic range of P. fa/cata, described in 1992 from dead cranberry
leaves in Massachusetts and New Jersey, is expanded with the discovery
of the fungus on dead leaves from two cranberry bogs in Wisconsin.
Key Words: cranberry microfungi, Pseudotracylla dentata, Pseudotracylla
falcata, pycnothyrium, Vaccinium macrocarpon
The pycnothyrial genus Pseudotracylla Sutton & Hodges (1976)
was based on a single collection of Euca/yptus citriodora Hook. leaves
from Brazil. Pseudotracylla dentata Sutton & Hodges, the type species,
is characterized by fusiform conidia and a dentate margin on the upper
pycnothyrial wall. Pseudotracylla remained monotypic until Carris (1992)
described P. falcata Carris from dead leaves and decayed fruit of
Vaccinium macrocarpon Ait. (cranberry) from Massachusetts and New
Jersey. Pseudotracylla falcata differs from P. dentata in having a smooth
margin on the upper pycnothyrial wall and distinctly falcate conidia. An
unusual type of conidiomatal development was described in P. falcata in
which a central, flattened, lobed cell expands outward, forming septa that
delimit individual cells. This single layer of cells eventually forms both the
upper and lower walls of the pycnothyrium. The limited amount of
material in the type collection of P. dentata precluded a precise description
of conidiomatal development in that species. Recently, P. dentata was
found on dead cranberry leaves from a commercial cranberry bog in
Massachusetts. The fungus was established in pure culture in order to
compare sporulation with that observed in P. fa/cata. Pycnothyria on
94
cranberry leaves were fixed and sectioned to compare conidiomatal
structure with the earlier report on P. falcata (Carris, 1992).
MATERIALS & METHODS
Leaves of V. macrocarpon were collected from piles of debris in
cultivated bogs in Massachusetts and Wisconsin during October and
November, 1990 and 1991. Leaf samples were stored in plastic bags at
5 C for up to three months. Production of conidia was stimulated by
placing leaves with pycnothyria in a moist chamber for one or more
weeks. Conidia were streaked onto Difco Bacto-agar and subcultures
made from single, germinating conidia. Isolates were grown in 9-cm diam
plastic Petri dishes on Difco corn meal agar (CMA) under artificial light and
temperature (20-25 C). Conidiomata were fixed and sectioned at 8 ym as
previously described (Carris, 1992). Cultures were deposited in ATCC,
CBS and IMI, and dried specimens were deposited in IMI and WSP.
RESULTS
Pseudotracylla dentata was found in leaf debris samples from one
of 24 cultivated cranberry beds surveyed in Massachusetts.
Pseudotracylla falcata also was present in this sample. Pycnothyria of one
or both species of Pseudotracylla were found on 70% of the leaves
examined. Pseudotracylla falcata was present on 56% of the colonized
leaves, P. dentata on 22%, and pycnothyria of both species were present
on the remaining 22% of the colonized leaves. Pseudotracylla falcata was
found in leaf debris samples from two of twelve cultivated cranberry beds
sampled in Wisconsin, expanding the known geographical range of this
fungus. Pseudotracylla dentata was not found in any of the Wisconsin
samples. Neither species has been found in cultivated cranberry bogs in
Oregon and Washington. The Massachusetts cranberry isolate of P.
dentata agrees with the published description (Sutton and Hodges, 1976)
and type material (IMI 196483k). The Wisconsin isolates of P. falcata
were indistinguishable from the holotype (WSP 69554) and other isolates
from Massachusetts and New Jersey. The reader is referred to Carris
(1992) for a description of this fungus on host tissue and in culture. A
description of P. dentata on host material and in culture follows.
Conidiomata on host tissue 48-132 ym diam, formed singly or
coalescing on lower leaf surface. Conidiomata superficial, attached to the
leaf by a single, narrow hypha penetrating the host epidermis. Conidioma
composed of septate upper wall connected to lower, membranous wall by
central columella (Fig. 1). Upper wall of conidioma (Figs. 2, 4) dark
reddish brown, composed of one central, globose-angular cell (=
columella) 6.2-8.8 x 5.3-7.9 ym diam; columella surrounded by 2-3
95
irregular rows of smaller globose-angular cells (= conidiogenous cells)
4.4-5.3 x 2.6-4.4 um; these cells merging into outwardly radiating, square
to rectangular cells (2.6-5.3 x 1.8-3.5 wm); terminating in a dentate
margin. Lower wall dark brown, membranous, composed of radial rows
of sparsely septate cells 1.8-3.5 wm diam (Fig. 3).
One-week-old colony on CMA 10-12 mm diam, with sparse,
hyaline, mostly submerged hyphae 2.7-6.3 wm diam, with numerous
clusters of dark cells submerged and on surface of agar (Fig. 5); clusters
formed on surface of agar developing radiating, white aerial hyphae (Fig.
6). Conidiogenous cells enteroblastic-phialidic, of two types: first type
hyaline, ampulliform to cylindrical, walls smooth, 6.3-13.5 x 2.6-4.1 ym,
with prominent, cylindrical collarettes 1.8-3.5 wm deep and 2.6-3.5 wm
wide, occasionally with percurrent proliferation resulting in annellations,
formed singly or on branched conidiophores (Figs. 8, 9); second type dark
brown, globose to ampulliform, becoming lobed, walls roughened, 5.3-8.8
x 4.4-7.9 ym, with inconspicuous collarettes, formed in clusters up to
850 um diam (Fig. 10). Conidiogenous cells developing into clusters of
lobed cells which form either irregular masses, or radiating shield-like
structures reminiscent of upper wall of pycnothyrium. Similar conidia
produced by both types of conidiogenous cells: hyaline, smooth, aseptate,
fusiform, apex tapered, base truncate, with inconspicuous gelatinous
appendage, 10.6-12.3 x 2.6-4.1 wm (Fig. 7), formed abundantly and
accumulating in gloeoid mass around conidiogenous cells, germinating
readily via single germ tube.
SPECIMENS EXAMINED: Pseudotracylla dentata, on leaves of Eucalyptus
citriodora, Brazil, Paulista, Pernambuco, 30-VI-1975 (HOLOTYPE: IMI
196483K); on leaves of Vaccinium macrocarpon, Massachusetts,
Plymouth County, South Carver, Wankinquoah Bog, 10-X-1990, F. Caruso
(WSP 69558; IMI 352172).
ADDITIONAL SPECIMENS EXAMINED: Pseudotracylla falcata, on leaves
and decayed fruit of Vaccinium macrocarpon, Wisconsin, Monroe County,
Potter Cranberry Company, 7-XI-1991, S. N. Jeffers (WSP 69586);
Russell Rezin & Sons, Inc., 7-XI-1991, S. N. Jeffers (WSP 69587).
Colonies of P. dentata are white and composed of hyaline
mycelium, whereas those of P. fa/cata are composed of dark brown
mycelium. Both species produce clusters of dark brown conidiogenous
cells, although the cells forming in cultures of P. dentata are mostly lobed,
and in P. falcata are globose. The hyaline conidiogenous cells with
prominent collarettes which were common in P. dentata cultures were not
observed in P. falcata. Growth rate of P. falcata colonies was
approximately twice that of P. dentata. When the two species were
96
i,
paired in culture, mycelium of P. fa/cata would eventually grow completely
around the slower growing colony of P. dentata without overgrowing
mycelium of the latter species.
DISCUSSION
Examination of the P. dentata isolates on cranberry leaves
substantiates the previous description of pycnothyrial structure in
Pseudotracylla which was based primarily on P. fa/cata (Carris, 1992). In
their original description of P. dentata, Sutton and Hodges (1976) had
described the upper pycnothyrial wall as sparsely septate and the lower
wall as composed of pseudoparenchyma. Hyphae composing the upper
wall of mature pycnothyria of P. dentata examined in this study were
regularly septate, whereas the lower wall was sparsely septate and
membranous. The membranous nature and sparse septation of the lower
wall (Fig. 4) suggest that it separated from the upper wall during an early
stage of pycnothyrial development when the upper wall was still
expanding.
The conidia produced in culture by the two types of
conidiogenous cells in P. dentata are indistinguishable. In contrast, the
conidia of P. falcata are only formed in culture from aggregations of dark,
thick-walled conidiogenous cells (Carris, 1992). The aggregates of
conidiogenous cells in Pseudotracylla species presumably represent an
early stage in pycnothyrial formation, although fully-developed pycnothyria
are rarely formed in agar culture. It is possible that pycnothyria develop
in response to contact with a specific type of surface, such as the leaf
cuticle. Pseudotracylla dentata is similar to Tubakia species in culture;
both form conidiogenous cells in aggregations and singly on hyphae
(Glawe and Crane, 1987). Munkvold and Neely (1991) questioned the
Figs. 1-10. Pseudotracylla dentata. 1. Two mature pycnothyria
grown together on leaf, longitudinal section (8 wm thick). Arrows indicate
columellae. X 800. 2. Upper wall of pycnothyrium, treated with NaOCl.
X 320. 3. Lower wall of pycnothyrium. X 640. 4. Upper wall of
developing pycnothyrium. X 640. 5. 11-da-old colony on corn meal
agar. X 5. 6. Clusters of dark conidiogenous cells produced on surface
and submerged in agar, 11-da-old colony on corn meal agar. X 19. 7.
Conidia produced in culture. Arrow indicates gelatinous sheath at base
of conidium. X 1350. 8, 9. Hyaline conidiogenous cells produced in
culture. X 1350. 10. Clusters of dark conidiogenous cells produced in
culture. X 1350.
98
placement of these types of fungi in the Coelomycetes since their
conidiomata are not true pycnidia or acervuli, and they grow like
Hyphomycetes in culture.
This report of P. dentata on cranberry leaf debris in
Massachusetts is notable in several regards. First, it is only the second
time the fungus has been found, and the first time it has been cultured.
Secondly, the occurrence of both known species of Pseudotracylla on
cranberry, at the same site, and indeed, on the same leaf, suggests some
type of host or habitat preference in the genus. However, as was
previously noted, the type collection of P. dentata is from Eucalyptus
leaves in Brazil. The existence of any similarity between the Brazil and
Massachusetts habitats and the Euca/yptus and Vaccinium hosts is not
obvious. Disparate host and geographical records such as this may be
more due to the inconspicuous nature of the fungi and our lack of
knowledge on them rather than to the actual distribution of species.
ACKNOWLEDGEMENTS
PPNS 0161, College of Agriculture and Home Economics
Research Center Project 0837, Washington State University, Pullman. |
thank Drs. Brenda Callan, Dean Glawe and Jack Rogers for manuscript
review, Drs. Frank Caruso and Steven Jeffers for providing leaf samples,
and Peter Gray for technical assistance.
LITERATURE CITED
Carris, L. M. 1992. Vaccinium fungi: Pseudotracylla falcata sp. nov.
Mycologia 84:534-540.
Glawe, D. A. and J. L. Crane. 1987. Illinois fungi. XIIl. Tubakia dryina.
Mycotaxon 29: 101-112.
Munkvold, G. P. and D. Neely. 1991. Development of 7ubakia dryina on
host tissue. Canad. J. Bot. 69: 1865-1871.
Sutton, B. C. and C. S. Hodges Jr. 1976. Eucalyptus microfungi:
Mycoleptodiscus species and Pseudotracylla gen. nov. Nova
Hedwigia 27: 693-700.
MY COTAXON
Volume L, pp. 99-105 January-March 1994
ACAULOSPORA EXCAVATA SP. NOV. - AN ENDOMYCORRHIZAL
FUNGUS FROM COTE D'IVOIRE
K. INGLEBY C. WALKER? & P.A. MASON'
"Institute of Terrestrial Ecology, Bush Estate,
Penicuik, Midlothian, Scotland, U.K. EH26 0QB
*Forestry Commission, The Forestry Authority,
Northern Research Station, Roslin, Midlothian,
Scotland, U.K. EH25 9SY
SUMMARY
A newly discovered endomycorrhizal fungus, Aucaulospora excavata, is
described and illustrated.
INTRODUCTION
As part of a study of spore populations of arbuscular mycorrhizal fungi in
Céte d'Ivoire (Wilson et a/. 1992), soil samples were collected from beneath
Terminalia superba Engl. & Diels and T. ivorensis A. Chev. in the Mopri
Forest Reserve, in October 1990. Among the spore types extracted was a
previously undescribed member of the genus Acaulospora Gerdemann and
Trappe. This fungus is described herein as Acaulospora excavata sp. nov.
MATERIALS AND METHODS
Spores were extracted from soil by sucrose centrifugation (Walker et a/. 1982)
and suspended in water for initial examination under a dissecting microscope
(x 6-50). Illumination was by incident light from an quartz-iodine fibre-optic
source with a colour temperature of 3200°K. Spore colour was described
from freshly extracted spores suspended in water in plastic Petri dishes and
illuminated by the same light source used for examining the spores. Colour
was determined by comparison with a standard colour chart (Anon. 1969), the
numbers following the colour descriptions refer to those given in the colour
chart. Colour matching of structures viewed with transmitted light under a
compound microscope was more difficult than with a dissecting microscope,
so for such observations colours (for example, of individual walls in the
description) were not precisely matched to a chart, and only generalised
colour descriptions are used (Walker et al. 1993).
100
Specimens were mounted in polyvinyl alcohol lacto-glycerol (PVLG) (Koske &
Tessier 1983) or distilled water (Spain 1990) with or without Melzer's reagent
(Morton 1986a) for further study under a compound microscope (x 150-1500)
with brightfield and Nomarski differential interference contrast illumination.
Wall structures were described using the standardized terminology and
murographs of Walker (1983) and Morton (1986b). So far, attempts to
establish this species in pot culture have failed.
ACAULOSPORA EXCAVATA Ingleby and Walker sp. nov. (Figs 1 & 2)
Sporae singillatim in terra enatae, lateraliter gestae in sacculo sporifero,
pallide ochraceae, ochraceae vel aurantiae, globosae vel subglobosae, 115-
200 x 100-165 wm. Sporarum tunicae 3 in turmis tribus: Turma A cum tunica
singula, colorata, 8-11 “wm crassa, foveis interdum leviter angularibus, 4-20 x
4-16 um in diam et 2-6 um profundis; turma B cum tunica singula, hyalina,
membranacea, 0.5-1.0 um crassa; turma C cum tunica singula, amorpha,
baccata, 1-2.5 ym crassa, in solutione Melzeri carnea vel purpurea.
SPORES borne singly in the soil laterally on the neck of a hyaline sporiferous
saccule that collapses after the spores mature; spores pale ochraceous (6) to
ochre (9) to orange (48); globose to subglobose, 115-200 x 100-165 ym.
SACCULE neck at point of attachment to the spore 9-19 um in diameter with
walls 1-2 um thick (Figs. 1A, 1B). Dimensions of saccule not determined.
WALL GROUP A ofa single, pale yellow, laminated wall, 8-11 um thick, not
reacting to Melzer's reagent. Outer surface ornamented by circular to
subcircular to elliptical pits, 4-20 x 4-16 zm in diameter and 2-6 um in depth
sometimes with a slightly angular outline (Figs. 1C, 1D). These pits appear
flat-bottomed with the basal area 3-15 x 3-12 um in diameter (Fig. 1C,
arrowed). Pits adjacent or up to 9 um apart.
Fig. 1. Light photomicrographs of spores of Aucaulospora excavata.
A. Saccule neck still attached to spore.
B. Saccule scar. Note the smaller pits immediately surrounding the scar.
C. Surface view of ornamented outer wall showing pit size, shape and
distribution. These pits are flat-bottomed and in some cases the basal
perimeter is in focus (arrowed).
D. A fractured spore showing the pits in cross-section.
E. A fractured spore showing all three walls. Note the beaded inner wall 3
which has expanded in PVLG and the area where the beaded deposition is
absent (arrowed).
1
10
102
SPORE WALL STRUCTURE of three walls (1-3) in three wall groups (A-C)
(see Murograph Fig. 2).
nN
Ww
Y
j
Y
]
g
Fig. 2 Murograph (after Walker 1983) of Acaulospora excavata.
WALL GROUP B of a single, hyaline, membranous wall, 0.5-1.0 um thick, not
reacting to Melzer's reagent (Fig. 15).
WALL GROUP C of a single 'beaded', amorphous wall, 1-2 um thick
(including beaded ornamentation on outer surface), which reacts strongly in
Melzer's reagent turning pink or purple within minutes of crushing the spores.
This wall is plastic in PVLG and, when crushed, may become up to 9 um
thick (Fig. 1E).
ETYMOLOGY: Latin excavata - referring to the appearance of the pits on the
outer wall.
DISTRIBUTION AND HABITAT: Known only from a single soil sample
collected from the Mopri Forest Reserve. The sampling area was one of
natural forest which had been manually cleared and burnt in 1989 before
replanting with T. ivorensis. This disturbance had resulted in marked
changes in soil chemistry with increases in pH (from 6.5 to 7.5), organic
matter and available nutrients; at the sample point where A. excavata was
found the pH was particularly high (8.2). Changes in vegetation also
occurred after the clearance. In addition to planting with T. ivorensis the plot
was rapidly colonised by a wide range of herbaceous species, most notably
Chromolaena odorata (L.) King & Robinson.
103
MYCORRHIZAL ASSOCIATIONS: Not known. Baiting out the species was
attempted by both single-spore and multi-spore isolation and open-pot culture
(Gilmore 1968) using Plantago lanceolata L. and Vigna unguiculata (L.) Walp.
as host plants. None of these attempts succeeded.
HOLOTYPE: Cote d'Ivoire: South West of Tissale - Mopri Forest Reserve
under Terminalia ivorensis, E; isotype OSC (Walker 1674. 30 X 90).
DISCUSSION
Spores of A. excavata possess a distinctly ornamented outer wall surface of
densely crowded, flat-bottomed pits. Six other species of Acaulospora have
been described so far which possess a distinctly pitted outer wall, namely
Acaulospora foveata Trappe and Janos (Janos & Trappe 1982), A. cavernata
Blaszkowski (Blaszkowski 1989), A. Jacunosa Morton (Morton 1986b), A.
paulinae Blaszkowski (Blaszkowski 1988), A. undulata Sieverding (Sieverding
1988) and A. scrobiculata Trappe (Trappe 1977).
Spores of A. foveata can be distinguished from A. excavata by their generally
larger size (185-310 x 215-350 tm) and colour (yellow/brown to red/brown to
black/brown). Spores of A. paulinae are smaller (60-95 um) and those of A.
scrobiculata are a paler colour (hyaline to olive to brown) while those of A.
undulata are both smaller (55-85 um) and are a paler colour (hyaline) than
those of A. excavata. We have not been able to examine spores of A.
cavernata , but from their description, they are similar in size and colour to
those of A. excavata.
From their descriptions, A. cavernata, A. lacunosa, A. paulinae and A.
scrobiculata have smaller pits than A. excavata (always < 6 um diam
compared with 4-20 x 4-16 um) and thinner outer walls (< 6 um thick
compared with 8-11 um). Pits of A. lacunosa, A. paulinae and A.
scrobiculata are also more widely spaced and irregularly shaped. Additionally
spores of A. cavernata are described as having a second wall in wall group A
and a coriaceous wall in wall group C while those of A. Jacunosa have a
second wall in wall group B. Spores of A. undulata possess an even thinner
ornamented wall (1-1.5 um thick) consisting of depressions separated by
ridges 1 um wide. They are also described as having an outer evanescent
wall, though examination of specimens from the type culture does not show
this phenomenon. Although A. excavata and A. foveata have similar sized
pits, the smaller spore size and more densely crowded pits of A. excavata
combine to give the surface of the outer wall a completely different
appearance to that of A. foveata.
Wall group C of A. excavata is described as a single beaded wall, though it
can appear as two walls in PVLG mounts. The depositions on the outer
surface of this amorphous wall clearly do not constitute a distinct wall, though
104
in places they are so crowded as to seemingly form a separate layer. In other
places, however, the beads may be absent, possibly due to detachment
during crushing (Figure 1E, arrowed).
ACKNOWLEDGEMENTS
This project was funded by the UK Overseas Development Administration and
the Commonwealth Development Corporation, and was contracted to ITE by
the International Forest Science Consultancy. Work using imported soil was
carried out under DAFS licence number IP/MISC/28/90 issued under the Plant
Health (Great Britain) order 1987. We wish to thank Dr J.M. Trappe, Oregon
State University, Department of Forest Science for preparing the latin
diagnosis and for his helpful review of the manuscript.
LITERATURE CITED
ANON, 1969. Flora of British fungi colour, identification chart. Royal
Botanic Garden, Edinburgh, HMSO.
BLASZKOWSKI, J. 1988. Three new vesicular-arbuscular mycorrhizal fun4i
(Endogonaceae) from Poland. Bull. Pol. Acad. Sci: Biol. Sci. 36: 271-275.
BLASZKOWSKI, J. 1989. Acaulospora cavernata (Endogonales) - a new
species from Poland with pitted spores. Crypt. Bot. 1: 204-207
GILMORE, A.E. 1968. Phycomycetous mycorrhizal organisms collected by
open-pot culture methods. Hilgardia 39: 87-105.
JANOS, D.P. & TRAPPE, J.M. 1982. New Acaulospora species from
America. Mycotaxon 15: 515-522.
JENKINS, W.R. 1964. A rapid centrifugal-flotation technique for
separating nematodes from soil. Plant Disease Report 48: 692.
KOSKE, R.E. & TESSIER, B. 1983. A convenient permanent slide mounting
medium. Mycological Society of America's Newsletter, 34(2): 59.
MORTON, J.B. 1986A. Effects of mountants and fixatives on wall structures
and Melzer's reaction in spores of two Acaulospora species
(Endogonaceae). Mycologia 78: 787- 794.
MORTON, J.B. 1986B. Three new species of Acaulospora (Endogonaceae)
from high aluminium, low pH soils in West Virginia. Mycologia 78:
641-648.
SIEVERDING, E. 1988. Two new species of vesicular arbuscular mycorrhizal
fungi in the Endogonaceae from tropical highlands of Africa. Agnew.
Botanik 62: 373-380.
SPAIN, J.L. 1990. Arguments for diagnoses based on unaltered wall
structures. Mycotoxon 38: 71-76.
TRAPPE, J.M. 1977. Three new Endogonaceae: G/omus constrictus,
Sclerocystis clavispora and Acaulospora scrobiculata. Mycotaxon 6:
359-366.
WALKER, C. 1983. Taxonomic concepts in the Endogonaceae: spore wall
characteristics in species descriptions. Mycotaxon 18: 443-455.
105
WALKER, C., MIZE, C.W. & McNABB, H.S. 1982. Populations of
of endogonaceous fungi at two locations in central lowa. Canadian Journal
of Botany 60: 2518-2529.
WALKER, C., GIANINAZZI-PEARSON, V. & MARION-ESPINASSE, H. 1993.
Scutellospora castanea, a newly described arbuscular mycorrhizal
species. Submitted to Cryptogamie.
WILSON, J., INGLEBY, K., MASON, P.A., IBRAHIM, K.D. & LAWSON, G.J.
1992. Long-term changes in VA mycorrhizal spore populations in
Terminalia plantations in Cote d'lvoire. In: Mycorrhizas in Ecosystems.
D.J. Read, D.H. Lewis, A.H. Fitter and |.J. Alexander (Eds.) C.A.B.
International, Wallingford, U.K.
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MY COTAXON
Volume L, pp. 107-113 January-March 1994
STUDIES ON KERATINOPHILIC FUNGI. II.
CHRYSOSPORIUM PILOSUM sp. nov.
J. GENE’, J. GUARRO’, K. ULFIG’, P. VIDAL' and J. CANO!
' Unitat de Microbiologia, Facultat de Medicina, Universitat Rovira i Virgili, 43201
Reus, Spain. ? Environmental Pollution Institute, 40832 Katowice, Poland.
ABSTRACT
Chrysosporium pilosum sp. nov., a keratinophilic hyphomycete isolated from river
sediments and soil, is described. The fungus is characterized by its restricted growth on
agar media at 25/37 °C and the production of conidia on deflected short protrusions, but
its most prominent feature is the presence of broad, simple, thick-walled, brownish,
sterile hyphae. The taxonomic position of the species is discussed.
RESUMEN
Se describe un nuevo hifomiceto keratinofilico, Chrysosporium pilosum sp. nov., a
partir de tres cepas aisladas de sedimentos fluviales y de suelo. Dicho hongo se
Caracteriza por presentar un crecimiento restringido en cultivo a 25/37 °C, por la
produccién de conidios a partir de cortas protuberancias, a menudo curvadas, y por la
existencia de hifas estériles de color pardo. Se discute la posicién taxondémica de esta
especie.
108
In our studies on keratinophilic fungi from different sources, many species of taxonomic
interest have been isolated. One of them, Chrysosporium pilosum sp. nov., is
described and illustrated below.
Chrysosporium pilosum Gené, Guarro et Ulfig sp. nov.
Etym.: referring to the sterile and brown hyphae.
Coloniae in agaro PYE ad 25 °C lente crescentes, elevatae, in medio convolutae,
Jrequenter in margine radiato-striatae, luteolae vel subbrunneae, floccosae vel coactae,
margo definitus, leviter lobulatus; reversum brunneoaurantiacum vel fuscum, 37 °C
crescit. Hyphae vegetativae hyalinae vel albolutae, ramosae, septatae, tenuitunicatae,
laeves, 1-2.5 um crassae; post 2-3 hebdomades hyphae steriles formantur, simplices,
brunneae et crassitunicatae, 3-5.5 ym crassae. Hyphae reticulatae nonnumquam
praesentes. Conidia terminalia et lateralia numerosa, oriunda ex brevibus protrusionibus
vel ramulis lateralibus, rectis vel flexuosis, nonnumquam Sessilia, Solitaria, luteola,
primum laevia et tenuitunicata, deinde verrucosa et crassitunicata, unicellularia,
globosa, subglobosa vel pyriformia, raro fusiformia, 4-6 x 3.5-5.5 um, cicatrice basilari
1,5-2 pm diam. Conidia intercalaria minus numerosa, Solitaria, luteola, primum laevia
et tenuitunicata, deinde verrucosa et crassitunicata, unicellularia, subglobosa,
doliiformia, nonnumquam unilateraliter inflata, 3.5-5.5 x 3-4 ym. Chlamydosporae
absentes. Species moderate keratinolytica. Reproductio sexualis ignota.
Holotypus: IMI 356294, ex sedimentis fluvialibus. \sotypus: FMR 4157.
On phytone-yeast extract agar (PYE) at 25 °C, colonies restricted, with a mean daily
radial spread of 0.4-0.7 mm, raised up to 6 mm, smooth or convoluted at the centre,
often radially folded at the periphery, yellowish-white to mustard-yellow, light brown
in age, consisting of a basal felt covered by cottony to floccose aerial mycelium, often
with droplets of colourless or light brown exudate, margin well-defined, slightly
lobulate; reverse brownish-orange or dark brown. Sporulating slowly. Hyphae hyaline
Fig. 1. Chrysosporium pilosum IMI 356294. a,b. Colonial morphology after 4 weeks
at 25 C (a. on PYE; b. on OMA). c-g. Fertile hyphae with terminal, lateral and
some intercalary conidia (c. bar=20um; d-f. bar=10um; g. bar=2pm). h.
Conidia (bar=1pm). i. Sterile hyphae (bar =20um).
109
110
Fig. 2. Chrysosporium pilosum |MI 356294. Fertile hyphae and conidia.
11]
to yellowish, branched, septate, smooth- and thin-walled, 1-2.5 »m wide; after 2-3
weeks (on all media tested) numerous, brown and thick-walled, sterile hyphae are
present, especially in the centre of the colony. These hyphae are simple, straight or
slightly wavy, sparsely septate, 3-5.5 um wide, bearing irregular, darker brown knobs.
Racquet hyphae occasionally present. Terminal and lateral conidia abundant, usually
born on deflected short protrusions or on straight or flexuose side branches, sometimes
sessile, solitary, hyaline to pale yellow, initially smooth- and thin-walled, becoming
thick-walled and usually coarsely verrucose at maturity, unicellular, globose, subglobose
to pyriform, in young cultures occasionally fusiform (IMI 356295), 4-6 x 3.5-5.5 um,
with basal scars 1.5-2 ym wide. Intercalary conidia often present (abundant in [MI
356295), solitary, pale yellow, at first smooth- and thin-walled, later becoming thick-
walled and often coarsely verrucose, unicellular, subglobose to barrel-shaped,
occasionally unilaterally inflated, 3.5-5.5 x 3-4 ym. Chlamydospores not observed.
Teleomorph unknown.
On oat-meal agar (OMA), colonies with a mean daily spread of 0.6-0.7 mm, raised (3-4
mm), smooth, slighly depressed at the centre, densely cottony, pale yellowish, becoming
light brown or brown at the centre due to dark hyphae, sometimes with colourless
exudate droplets, margin defined, irregularly lobulate, slightly fimbriate; reverse
reddish-orange or olive-brown, with the yellowish pigment diffusing into the medium.
Sporulation abundant. With the strain IMI 356790, we obtained occasionally a structure
similar to a hairy pseudoascomata but it was sterile.
The fungus grows at 37 °C, with mean daily radial spread of 0.3-0.4 and 0.6-0.7 mm
on PYE and OMA, respectively. Colonies on PYE yellowish-white, raised up to 3-4
mm, slightly convoluted, with a basal felt covered by cottony to floccose aerial
mycelium; margin defined, irregular; reverse colourless or light brown. On OMA,
colonies yellowish-white, floccose and raised up to 2-4 mm at the centre, white,
funiculose and radially folded at the periphery; margin defined, lobulate; reverse pale
green, with the yellow pigment diffusing into the medium.
Resistant to cycloheximide 0.2 %. Slightly keratinolytic.
Material examined: IMI 356294 (=FMR 4157), from river sediments of the River Fluvia, Catalonia, Spain,
16 September 1991; IMI 356295 (=FMR 3972, RV 24220), from soil, Somalia, 14 March 1968; IMI 356790
(=FMR 4345), from forest soil, Guanajay, Pinar del Rio, Cuba, 22 March 1992.
TZ
Fig. 3. Chrysosporium pilosum \MI 356294. Sterile hyphae.
The main characteristic of both isolates of C. pilosum, unique for this genus, is the
presence of brown, thick-walled hyphae, suggesting an ascomycete teleomorph.
However, no ascomata could be obtained under a variety of growth conditions (different
temperatures and media) and when the three strains were crossed with each other.
Species with a similar characteristic are Onychocola canadensis Sigler et Congly
(1990) and Malbranchea filamentosa Sigler et Carmichael (Sigler et al., 1982). Both
species have arthroconidia and in neither case did a teleomorph develop. Some other
species of Chrysosporium, such as C. merdarium (Link) Carmichael (1962), the C.
anamorph of Amaurascopsis perforatus Guarro et al. (1992), the C. anamorph of
Renispora flavissima Sigler et al. (1979), and C. vallenarense Oorschot et Piontelli
(1985), resemble C. pilosum in having yellow pigmentation of the colonies as well as
globose to obovoid and coarsely verrucose conidia. However, C. merdarium and C.
vallenarense have a maximum temperature for growth of about 30 °C. In addition, the
growth rate of the former is faster at 20 °C, and the conidia in the latter are often
iS
produced on sympodial branches. The C. anamorph of R. flavissima produces larger
(5-12 x 4-12 pm), spiny or prominently tuberculate conidia, whereas that of A.
perforatus has a faster growth at 37 °C, with a Magenta-coloured colony reverse on
OMA, producing a teleomorph with peridial hyphae quite different from the sterile
hyphae observed in C. pilosum.
ACKNOWLEDGEMENTS
This work was supported by grant n° 92/0785 from FIS from the Ministerio de Sanidad
y Consumo, Spain. The authors are indebted to Dr. E. Descals (C.S.1.C.), Dr. C.S.
Tan and Prof. W. Gams (CBS, Baarn) for helpful comments and to Dr. J.M. Escola
(Universitat Rovira i Virgili, Tarragona) for reviewing the Latin diagnosis. We also
express our thanks to Prof. Ch. de Vroey (Institute of Tropical Medicine ‘Prince
Leopold’, Antwerp, Belgium) who supplied an isolate.
REFERENCES
Carmichael J.W. 1962. Chrysosporium and some other aleuriosporic Hyphomycetes.
Can. J. Bot. 40: 1137-1173.
Guarro J., J. Gené and Ch. de Vroey. 1992. Amaurascopsis, a new genus of
Eurotiales. Mycotaxon 45: 171-178.
Oorschot C.A.N. van and E. Piontelli. 1985. Chrysosporium vallenarense spec. nov.
Persoonia 12: 487-488.
Sigler L. and H. Congly. 1990. Toenail infection caused by Onychocola canadensis
gen. et sp. nov. J. Med. Vet. Mycol. 28: 405-417.
Sigler L., P.K. Gaur, R.W. Lichtwardt and J.W. Carmichael. 1979. Renispora
flavissima, a new gymnoascaceous fungus with tuberculate Chrysosporium conidia.
Mycotaxon 10: 133-141.
Sigler L., J. Lacey and J.W. Carmichael. 1982. Two new species of Malbranchea.
Mycotaxon 15: 465-471.
MY COTAXON
Volume L, pp. 115-122 January-March 1994
A NEW SPECIES OF DELITSCHIA FROM WEST AFRICA
FERNANDO PEL AEZ
Merck Sharp & Dohme de Espana, S.A.
Josefa Valcarcel, 38
28027 Madrid, Spain
JON D. POLISHOOK
Merck Research Laboratories
P.O. Box 2000, Rahway, New Jersey 07065, USA
MARTI VALLDOSERA AND JOSEP GUARRO
Unitat de Microbiologia, Facultat de Medicina
Universitat Rovira | Virgili, 43201 Reus, Tarragona, Spain
Summary
A new species of Delitschia growing in culture is
described and illustrated. The fungus was isolated from
dung of a dassie collected in Namibia. It is characterized by
having asci each with 64 ascospores. Also, each ascospore
has a transverse and non-constricted septum, with a
longitudinal germ slit.
Introduction
Species included in the genus Delitschia Auersw. are
fimicolous loculoascomycetes, characterized by bitunicate asci and
dark brown to black, two-celled ascospores. The septum is median,
transverse or oblique, and both cells of the ascospore have an
elongated germ slit. A hyaline gelatinous sheath surrounds each
116
ascospore. The genus was last reviewed in 1975 by Luck-Allen and
Cain (1), who recorded 46 species. Later, six more species were
added (2,3). Of these 52 total species, only 6 produce asci with
more than 8 ascospores, with the remainder having asci with 8 or
less ascospores.
In this paper we describe a new species of Delitschia with
approximately 64 ascospores per ascus isolated from dung of
dassie (Procavia sp.) collected in Namibia. A key to the species of
this genus having more than eight ascospores is also presented.
Materials and Methods
Isolation of the culture: 0.5 g of dried dung was ground and
rigorously washed with water and air for 1 hour in a device with
four screens decreasing in pore size, similar to the one described
by Parkinson & Williams for the isolation of soil fungi (4). The
particles retained in the smallest size screen (90-105 um) were
plated onto VDY medium, prepared according to Papavizas & Davey
(5). The growing colonies were transfered to plates of potato
dextrose and oatmeal agar for identification. Mature perithecia
were observed after 2-3 weeks growing at 249°C under fluorescent
light (near UV) on a 12 hour light/dark cycle.
In the descriptions capitalized color names in parentheses are
from Ridgway (6). Cornmeal agar and V-8 juice agar were prepared
as media M-7 and M-29, respectively, of Stevens (7). Potato
dextrose agar and oatmeal agar were prepared media from Difco
Laboratories.
Fig. 1. Delitschia confertaspora. A. Perithecium x 100. B. Peridium
in surface view x 1200. C. Mature ascus x 320. D . Mature and
immature asci, pSeudoparaphyses and ascospores x 100. E. Upper
portion of an ascus showing the double wall, the apical structure
and mature ascospores x 500. F. Mature ascospore showing the
transverse septum and the gelatinous sheath expanding about the
ends of the spore x 1250. G. Mature ascospore showing the
longitudinal germ slit x 1200. H. Mature ascospores with gelatinous
sheath x 500.
Dy
118
Description of the species
Delitschia confertaspora Pelaez, Polishook, Valldosera and
Guarro, sp. nov. Figs. 1-2
Peritheciis subglobosis vel piriformibus sparsis, brunneis vel
atrobrunneis, 400-625 x 350-625 tum, pilis hyalinis vel flavo-
brunneis, septatis, 2-4 um diam, collo papilliformi vel cylindraceo,
65-150 x 80-130 um; peridio membranaceo vel aliquantum
coriaceo, atrobrunneo, opaco; cellulis externis irregulariter
angulatis vel epidermoideis. Ascis 64-sporis, cylindraceis vel
Clavatis, 250-365 x 37-57 tm, superne late rotundatis, brevi-
Stipitatis; pseudoparaphysibur numerosis, hyalinis, filiformibus,
0.8-4 um diam, septatis. Ascosporis multiseriatis, brunneis vel
atrobrunneis, opacis, ellipsoideis, 23-25 x 9-11 um, transverse
uniseptatis, leviter constrictis vel nonconstrictis, segmentis
ascosporarum maturiS sape sejuntis, strato mucoso hyalino
involutis, stria germinal longitudinaliter prolata.
TYPO: In fimo ex dassie, Mirabeb Hills, Namibia, 23 Sept. 1990.
Teges culta sicca IMI 350,692 (=ATCC 74209).
ETYMOLOGY: Latin, conferta = crowded or dense, spora = seed,
referring to the arrangement and quantity of spores in the ascus.
Perithecia dark brown to black, subglobose to pyriform, 400-
625 x 350-625 um; irregularly covered with hyaline to pale brown,
septate, mostly unbranched hairs, 2-4 um in diam; scattered,
solitary to aggregated, superficial; ascospores, when mature,
extruded as black, slimy mass; neck dark brown, papillate to short
cylindrical, 65-150 x 80-130 um; peridium membranaceous to
rather coriaceous, dark brown, opaque, consisting of irregularly,
thick-walled cells, textura angularis to epidermoidea.
Asci usually 64-spored, bitunicate, cylindrical to clavate,
broadly rounded above, tapering below into a short stipe, 250-365
X77 tm:
Pseudoparaphyses numerous, hyaline, filamentous, septate,
0.8-4 um in diam. Ascospores multiseriate, brown to dark brown,
ellipsoidal, broadly rounded at the ends, 23-25 x 9-11 um,
transversely septate, 2-celled, unconstricted or only slightly
constricted at the median septum and non separable at maturity,
119
Fig. 2. Delitschia confertaspora. A. Perithecium. B. Ascus.
C. Ascospores, two of them showing the gelatinous sheath.
Bar: A = 140 um; B = 35 um; C = 10 um.
120
each cell with a longitudinal germ slit, surrounded by an uneven
hyaline gelatinous layer expanding up to 12 um around the spore,
wider on the ends of the spore than near the septum, and only
visible in fresh preparations from young cultures.
CULTURAL CHARACTERISTICS: On cornmeal agar colonies
growing moderately slow and submerged, attaining a diameter of
34 mm in 14 days at 259C and 50% RH, margin uneven, surface
appressed, sparse with margin dissected, culture mat near
inoculum light brown (Clay Color, Cacao Brown) fading to colorless
at margin, immature perithecia present.
On potato dextrose agar attaining a diameter of 41 mm, aerial
mycelium cottony in colony center to a flat, mostly appressed
margin, radially sulcate, margin uneven, color light orange-yellow
(Light Ochraceous Buff, Ochraceous Buff) with a light gray (Dawn
Gray) near the margin, reverse reddish brown (Burnt Sienna,
Sandfords Brown), soluble pigment present, perithecia absent.
On oatmeal agar attaining a diameter of 41 mm, culture mat
radially sulcate with some media buckling, mat near inoculum light
brown (Light Ochraceous Buff, Ochraceous Buff), margin darker
brown (Pecan Brown, Cinnamon), light yellow-brown exudate
present at inoculum point, mature perithecia present.
On V-8 juice agar attaining a diameter of 44 mm, culture mat
appressed or slightly raised, floccose to cottony at inoculum
sometimes radiating to margin, color light brown (Flesh Ocher,
Apricot Buff) with some sections with a gray tint (Hathi Gray),
mature perithecia present.
HABITAT: On dung of dassie (Procavia sp.).
SPECIMEN EXAMINED: Mirabeb Hills, Namibia, Africa. 23
September 1990. Dried culture mat IMI 350,692 (=ATCC 74209).
Discussion
Delitschia confertaspora is characterized by having 64-spored
asci, each ascospore with a longitudinal germ slit and a transverse
and unconstricted septum. Only two other species of Delitschia
with 64 ascospores per ascus have been previously described (2):
D. dochmiophragmia and D. spiralirima. The species described here
is easily distinguishable from these two.
The ascospores of D. spiralirima have a distinct spiral-shaped
124
germ slit and are significantly larger in size (51-57 x 24-29 um)
than the ascospores of D. confertaspora, which have a longitudinal
germ slit, and also the asci are much larger in D. spiralirima (450-
600 x 130-150 um).
In D. dochmiophragmia the ascospores have an oblique and
constricted septum with each cell separating at maturity. D.
confertaspora, however, has ascospores with a transverse septum,
unconstricted or only very slightly constricted, and with the cells
remaining intact at maturity. In addition, the ascospores of D.
confertaspora are slightly smaller in size than D. dochmiophragmia
(27-39 x 12-15 um), and with the ends more broadly rounded than
in this last species.
Finally, the gelatinous sheath that surrounds each ascospore of
D.confertaspora, expanding about the ends, is _ clearly
distinguishable from the other two species mentioned. However, it
resembles the sheath of D. mesostenospora, although this species
has eight-spored asci.
The following key discriminates among the described species
of Delitschia with more than eight ascospores per ascus.
SON ulti OTA SCOS DOES eee eric tn nye ee deer tee en seen rates Nt oa 2
ASCII One mat d OvaSCOSDOIOS tits. ste cs te. eer recent fot tdasaantcede dares 3
—_
2. Ascospores with septum transverse; 56-62 x 18-21 um
Re meen oh Re en he ote NL A ah aaetech eh te Tad Cond D. polyspora Griff.
2. Ascospores with septum oblique; 36-54 x 12-15 um
ct ch EAE RES OEP Tn D. sexdecimspora Jeng, Luck-Allen & Cain
PPASEM WITT See ASCliic ont ac soe urecass D. ionthada Jeng, Luck-Allen & Cain
SRAcGEWMOrouial 21a SCOSDONCS male yar cuticle Cee aan es aie, 4
APASCIMIMIN OG: ASCOSDOLC Giamce pe ssudenn cath teat teach: | Poem nin, deh Plena BS es 5
4. Asci with 256 ascospores ................. D. myriaspora Breton & Faurel
5. Ascospores oblong with spiral shaped germ slit
DR cero Batra. RMON milena eae RAY Rs D. spiralirima Jeng, Luck-Allen & Cain
5. Ascospores ellipsoidal with longitudinal slits ................cceeeceeeeeee ees 6
6. Ascospores obliquely septate
EU Re Oe eee tere EAU ORRE Penn rerrnian D. dochmiophragma Jeng, Luck-Allen & Cain
6. Ascospores transversely septate ............... D. confertaspora sp. nov.
122
Acknowledgments
The authors would like to thank Dr. D. Jose Maria Barrasa,
University of Alcala de Henares, Madrid, Spain, for prepublication
review of the manuscript.
Literature
1. Luck-Allen, E.R. and Cain, R.F. 1975. Additions to the genus
Delitschia. Can. J. Bot. 53: 1827-1887.
2. Jeng, R.S., Luck-Allen, E.R. and Cain, R.F. 1977. New species and
new records of Delitschia from Venezuela. Can. J. Bot. 55: 383-392.
3. Furuya, K. and Udagawa, S. 1976 . Coprophilous pyrenomycetes
from Japan IV. Trans. mycol. Soc. Japan17: 248-261.
4. Parkinson, D. and Williams, S.T. 1961. A method for isolating
fungi from soil microhabitats. Plant and Soil13: 347-355.
5. Papavizas, G.C. and Davey, C.B. 1959. Evaluation of various
media and antimicrobial agents for isolation of soil fungi. Soil
Science 88: 112-117.
6. Ridgway, R. 1912. Color standards and color nomenclature.
Publ. by the author. Washington, D.C. 43 p. + 53 pl.
7. Stevens, R. B. 1981. Mycology Guidebook. University of
Washington Press. Seattle and London. 712 p.
MOY COWAXON
A NEW SPECIES OF MARAVALIA FROM INDIA
G. BAGYANARAYANA AND E. JOHN RAVINDER
Botany Department, P.G. College of Science
Saifabad, Hyderabad - 500 004 (A.P.) India
The genus Maravalia Arth. is characteristic in the
possession of subepidermal, erumpent Celia. with
pedicillate, one celled, thin walled, non resting
teliospores found successively on laterally free basidio-
genous cells (Cummins & Hiratsuka, 1983). SO wet ares L
species of Maravalia are reported (Ono, 1984) on members
belonging to various angiospermic families nie a a
Acanthaceae, Erythroxylaceae, Euphorbiaceae, Leguminosae,
Periplocaceae, Rubiaceae, Sapotaceae and Verbenaceae.
Recently, the authors came across a hitherto undescribed
species of Maravalia on Tylophora sp. (Asclepiadaceae) and
the same is reported here.
Maravalia ramacharii John Ravinder & Bagyanarayana_ sp.
NOV.) VRaG 3 1)
Spermagoniis. et aeciis ignotis.
Urediniis hypophylliis, Ssparsis, raro aggregatis,
sub-epidermalis, erumpentis, pulverulentis, pallide
aurantiaco flavescentis, 0O.7 - 1 mm crassa; uredinio-
SHorigs,..) Dedredllatic i Nise i 27 kl a be Un, OVaATLS.,
globosis vel ellipsoideis, membrana echinulata, hyalina,
2 um crassa, poris germinativis ignotiis.
Teliis hypophylliis, sparsis vel aggregatis 0.3- 0.6:
crassa, subepidermalis, erumpentis, teliosporiis 31 - 45 x
9 - 12.4 um, oblongo ellipsoideis, membrana 1.5..um crassa;
pedicillatis, pedicello hyalinis, usque ad 21 um) longis.
Molotyous: “i Inv foliis vivis \Tylophorae’sp.:,):Decw Lo,
Narsapur (A.P.), India, John Ravinder, HCIO; Isotype PUR.
124
Etymology : In honour of late Prof. P..Ramachar, a reputed
uredinologist of India.
Spermogonia and aecia not known. Uredinia hypo-
phyllous, scattered, rarely aggregated, subepidermal,
erumpent, ~pulverulent, . light orange yellow,;, 0.7 — 31 amma
diam; urediniospores pedicillate, 18 - 27 x 12.4 - 18 um,
ovate, globose to ellipsoid, orange yellow when fresh,
wall echinulate, hyaline, 2 um thick, germ pores absent.
Telia hypophyllous, scattered or aggregated in small
clusters; 0O.3$ — 0.6 > diam.; subepidermal, erumpent; telio-—
spores (G1 0-45.19" — 12. 47um: s0blong = eliapsoid 7 convenes
yellowish-orange’) when fresh, wall 1:5 um thick, hyaline,
smooth; (pedicrl late, pedicel hyaline, upto. zi. um long,
As per Ono's (1984) recent monograph on Maravalia,
host genera belonging to Asclepiadaceae are not infected
by species of Maravalia. However, Cryptostegia and
Pentopetia belonging to Periplocaceae, a closely related
family to Asclepladaceae . are infected by Maravalia
cryptostegiae (Cumm., ) Ono and Maravalia species.
M. ramacharii does not bear any morphological resemblance
to the Maravalia species reported on Periplocaceae.
The authors express their grateful thanks ‘to Prof:
Joe F. Hennen, Director, Arthur Herbarium and Head, Dept.
of "Botany ""S" Plants Pathology, Purdue™ University, =for
reviewing the manuscript and for helpful suggestions.
Fig. 1 Maravalia ramacharii sp. nov. (A) T.S. through the
sorus showing teliospores (B) A pedicillate teliospore
(stained) withicotton blue) (C) Am echinulate uréediniospore.
125
REFERENCES
Commans, “G.B.s.& Hiratsuka, Y., 1983. Illustrated genera of
rust fungi. Burgess Publishing Company, Minneapolis,
WrotswNc
OnGra).. ULoSe., A monograph of Maravalia (Uredinales).
Mycologia. 761(5):i¢ 692-911,
1; Vook
w
- +)
MY COTAXON
Volume L, pp. 127-129 January-March 1994
A NEW SPECIES OF UROMYCES ON OCIMUM FROM INDIA
G. BAGYANARAYANA AND E. JOHN RAVINDER
Botany Department, P.G. College of Science
Saifabad, Hyderabad - 500 004 (A.P.) INDIA
The family Labiatae (Lamiaceae) is known to have 3500
species distributed over 180 genera (Willis, 1982 Rev. by
Airy Shaw). Nearly 55 of these genera consisting of 325
species are reported from India (Hooker, 1985; Gamble,
1957; Cooke, 1958). Of the so many genera Ocimum of the
subfamily Ocimoideae is one of the most common genus
SOCCUBINGw ain S-wilde sas) well. as cultivated” jconditions.
Regent ly ,sauningy a study of the. rust: fungus flora’ vot
Andhra Pradesh, the authors have coliected rust infected
plants of Ocimum sp. from Mannanore forest, Mahabubnagar
Dist; (A Pa) & India. A. “critical (microscopic study
revealed an undescribed species of Uromyces and the same
is described here.
Uromyces ramacharii John Ravinder & Bagyanarayana
Speuneow. CPigns 2)
Spermagoniis et aeciis ignotis
Urediniis epiphylliis, subepidermalis, erumpentis,
pulverulentis, Ccinnamomeo' brunneis, 1-1.5 mm crassa;
uredainiosporius pedicillatis,) 16-23. x) 1525 ——.- lé.6Gieum,
globosis, .subglobosis, ovatis vel ellipsoideis, membrana
1.5 - 2 um crassa, pallide flavescentis vel hyalinis,
echinulatis, poris germinativis 2, equatorialibus.
Teliis amphigeniis, densus hypophylliis, atro-brunneis,
Sparsis vel aggregatis, subepidermalis, erumpentis,
pudverittentis:). 1 G32 min °crassa; “teliosporiis 21) —. 32) x
ie26) = 21 ums “qlobsis,, subglobosis’ vel ovatis>: membrana
lateralis 3.5-5.4 um crassa, raro apicalis, 6.3 um crassa,
verrucosis; pedicillatis, pedicello 15.5 - 45 um longis,
hyalinis.
128
Holotypus : In foliis vivis Ocimum sp. (Labiatae),
Mannanore forest, Mahabubnagar Dist. (A.P.), India Dec.
1990, Jagadeeswar and John Ravinder. HCIO. Isotype PUR.
Etymology : In honour of my (G.B.) revered teacher late
Prof. P. Ramachar, a reputed Uredionologist of India.
Spermagonia and accia not present. Uredinia
epiphyllous, subepidermal, erumpent, pulverulent, cinnamon
brown,” =) 125° "mm ‘diam; urediniospores pedictilate:
16) — 23; x% ofS.5 = 11656 (um globose, “subgiopose. “ovate,
ellipsoid, wall 1.5 - 2 um thick, pale yellow to hyaline,
echinulate, germpores 2, equatorial.
Telia amphigenous, densely hypophyllous, blackish brown,
scattered or aggregated, subepidermal, erumpent,
pulverulent, 1:=2;mm diam, .teliospores'21°-+'32x18.6 - 21
um, globose, subglobose, ovate, wall 3.5 - 5.4 um thick
laterally, . few spores, .are: thick apically, upto '6.3 um,
chestnut brown, verrucose, pedicillate, pedicel 15.5 - 45
um long, hyaline.
The members of the family Labiatae are known to be
parasitized by several rust fungi particularly the species
belonging to the genus Puccinia (Sydow & Sydow, 1904;
Arthur, 1934; Savulescu, 1953; Gaumann;*’ 1959; Wilson ©
Henderson 1966). However, only 4 species of Uromyces
viz., U.prunellae Schneid., U.lamii Kom, U.ocimi Hansford
and U.orthosiphoniis Ramakri. & Srinivasan are reported.
Of this Savulescu (1953) has listed U.prunellae as a
synonym of U.valerianae (Schum.) Fuckel, a species known to
parasitize, Valeriana of
the family Valerianaceae
but not Labiatae. With
regard to U.lamii, Sydow
& Sydow (1910) treated
this as an excluded
species because the_ so
called host Lamium
amplexicaule was found to
be a species of
Euphorbia. U.ramacharii
is comparable with U.ocimi
and U.orthosiphonis' the
two near species of
Uromyces on Labiatae.
U.ocimi differs from
U.ramacharii in i ihaving
only caulicolous telia,
longer teliospores with a
prominent apical thick-
ness upto 9 um and a
Fig. 1 Uromyces ramacharii sp. nov. (A) Echinulate
urediniospores (B) Pedicillate teliospores.
129
fairly long (160 um) persistent pedicels. U.ramacharii
differs from U.orthospihonis in lacking peripheral hyphoid
paraphyses in the uredinia, comparatively smaller
uridiniospores, and in having narrow but longer
teliospores.
The authors express their grateful thanks to Prof.
Joe F. Hennen, Director, Arthur Herbarium and Head, Dept.
of Botany and Plurnt Pathology, Purdue University, for his
kind help in reviewing the manuscript.
REFERENCES
1 Arthur, | C.J.) 1934. (Manuals of ithe irustsin “United
States and Canada. Purdue Research Foundation,
Lafayette, Indiana.
AE Cooke, T. 1958. The flora of the Presidency of Bombay
Vor. if, Botanical Survey :of India, ;Calcutta.
SN Gamble, J.S. 1957. Flora of the Presidency of Madras
Vor. Ii.. Botanical Survey ‘of India, Calcutta.
ay Gaumann, Es 1959. Die Rostpilze Mitteleuropas.
Buchdruckeres Buchter’ 6’ Co”. Bern,
Ss Hooker, J.D. 1875. The flora of British India, Reeve &
Com bOndon
6. Savulescu, °'T., 1953.0 Monografia. \Uredinalelor , 11.
Editura Academei Rupublicii Populare Romane.
Ais Sydow, P. & Sydow, H. 1910. Monographia Uredinearum.
VoL. I1,: bipsiae,, Fratres Borntraeger.
SB. Wario seer on ROW eg DV AEP V : MohaWay al 6 Kost) bow
dictionary of the flowering plants and ferns. Indian
reprint by International Distributors, Dehradun.
oc Wilson, M. & Henderson, D.M. 1966. British Rust Fungi
University Press, Cambridge.
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MY COTAXON
Volume L, pp. 131-138 January-March 1994
AMANITA WESTII—TAXONOMY AND DISTRIBUTION.
A RARE SPECIES FROM STATES BORDERING
THE GULF OF MEXICO
Rodham E. Tulloss David P. Lewis
P. O. Box 57 455 Virginia Lane
Roosevelt, New Jersey 08555-0057 Vidor, Texas 77662
Summary
Amanita westii was described from Florida oak woods. Collections
from Mississippi and eastern Texas extend its known range and allow
for an improved description.
During his years in Florida, William A. Murrill was a prolific collector of fungi and
described over 650 species of agarics and boletes as new. A number of these taxa have
not been reported since he described them 40 or more years ago. One such species, the
strikingly beautiful Amanita westii, was described by Murrill (1944) from a collection
made by Murrill’s associate Erdman West in Alachua County in 1938. The vegetation
of the Gainesville, Florida area, where Murrill resided, is similar to that of the Gulf
Coast region, and it may be surmised that the mycota would also be similar. It is from
the Gulf Coast region that we report new collections of A. westii.
This unusual, large Amanita was rediscovered by Lewis during a 1987 collecting
trip in eastern Texas. During the 1991 Gulf States Mycological Society summer foray,
Ms. Toby Feibelman made a collection of A. westii in southern Mississippi. The latter
collection included both mature specimens and ‘‘buttons.’’ Lewis studied both of these
collections when fresh and photographed them. The holotype of A. westii has been
studied previously by Bas (1969) and Jenkins (1979).
Methods and terminology follow those of Tulloss et al. (1992) and Tulloss (1993).
We cite specimens deposited in the following herbaria:
F - Herbarium, Field Museum of Natural History, Chicago, Illinois, U.S.A.
FLAS - Herbarium, University of Florida, Gainesville, U.S.A.
L - Rijksherbarium, Leiden, the Netherlands.
RET - personal herbarium of R. E. Tulloss.
Color codes of the form ‘‘7.5YR 8/6’’ are from (Munsell Color, 1975). Color
names in italics with first letters capitalized (e.g., Hair Brown) are from (Ridgway,
1912).
132
D.C. Tulloss del.
Fig. 1. Amanita westii (Lewis 4074) habit (x0.7).
133
AMANITA WESTI/ (Murr.) Murr. 1944 [1945]. Proc. Florida Acad. Sci. 7: 127.
=Venenarius westii Murr. 1944 [1945]. Proc. Florida Acad. Sci. 7: 115.
Illus.: Bas. 1969. Persoonia 5: 485, Figs. 244-246 [exsiccatum from holotype,
universal veil elements, spores].
Amanita westii (Figs. 1-2) has a pileus densely set with pyramidal warts that are
brown on the surface and white in the interior (at least in young material). The
basidiocarp is prone to reddish bruising and staining similar to that exhibited by A.
rubescens Pers. and has a slight odor of anise. The stipe has a radicating bulb. Dried
specimens take on a deep wine-red color somewhat like that of exsiccata of Lepiota
americana Peck.
PILEUS: 70 - 135+ mm wide (largest pileus measured not fully expanded), convex
to plano-convex, eventually planar, subviscid to dry, pale reddish brown (Murmill) to
pale chocolate brown (Murrill), cracks between scales 10R 9/1 (near Pallid Vinaceous
Drab), 10R 6/2 under warts, drying a dark wine color reminiscent of dried basidiocarps
of Lepiota americana; context up to 20 mm thick at the stipe, white quickly bruising
reddish brown or 7.5R 4/4 (near Prussian Red) changing to 10R 2/2 (near Light Seal
Brown), eventually blackish, the color of cocoa powder in exsiccata; margin nonstriate,
appendiculate with material hanging down 3 - 4 mm; universal veil as large warts,
pyramidal to somewhat flattened (in the latter case often with a small central pimple), 2
- 4 mm high, with irregularly polygonal bases 3 - 5 x 8 - 10 mm, largest near pileus
margin, densest over disk, 10R 2/2 (near Light Seal Brown), detersile.
LAMELLAE: adnexed by a line to stipe apex at first, then seceding, sometimes
remote at maturity, rather crowded to close to subdistant, white at first, then gray white,
finally reddish brown or 10R 6/2, drying reddish brown to dark brown, 11+ mm broad,
with edge thick and flocculose to fimbriate to almost crenate; lamellulae short, in one
or two tiers, with the shortest subtruncate to somewhat rounded subattenuate, with the
longest not distinguishable in exsiccata (notes from fresh material lacking).
STIPE: 70 - 155 x 15 - 35 mm, more or less cylindric or constricted at about the
mid-point, covered at first with floccose-fibrillose material which adheres to the fin-
gers, densely floccose to floccose-subfelted in apical region below partial veil, becom-
ing undecorated, dry, originally whitish (sometimes becoming grayish) near apex,
otherwise reddish brown or a very dark brown or pale reddish brown or 10R 6/2, con-
colorous with pileus in exsiccata; context white, with color changes on cutting or bruis-
ing as in pileus, solid; bulb obovoid-napiform to napiform to turbinate to broadly
fusiform, radicating, occasionally abrupt to subabrupt, up to 40 - 45* x 30 - 50 mm,
becoming colored like other parts; partial veil apical, subfelted to felted-
submembranous, fragile, detersile or adhering to edges of lamellae, white at first, then
pale reddish brown or 10R 6/2, striate above, underside densely flocculose; universal
veil as fibrillose material (occasionally vague warts or felted patches) at stipe base and
on upper half of bulb in “‘button’’ stage, detersile, colored as on pileus.
Odor faintly of anise (Murrill). Taste sweet and nutty at first, becoming slightly
astringent (Murrill).
134
D.C. Tulloss del.
Fig. 2. Amanita westii (Feibelman 1165) habit, ““button’’ specimens (x1.0).
135
MACROCHEMICAL TESTS: none recorded.
PILEIPELLIS: In mature material: 50 - 65 um thick, orange-brown in 3% KOH,
with surface extensively gelatinized and depigmented; filamentous, undifferentiated hy-
phae 1.0 - 9.8 uum wide, subradially oriented, tightly interwoven, branching; vascular
hyphae 5.0 - 17.5 um wide, branching, most easily seen in scalp. In ‘‘button’’: 45 - 60
um thick, ungelatinized, orange-brown to yellow-brown in 3% KOH. PILEUS CON-
TEXT: cell walls hyaline to yellow-brown to orange-brown in 3% KOH, with those
having thickest walls among (but not exclusively) the most strongly pigmented; fila-
mentous, undifferentiated hyphae 2.2 - 9.2 um wide, branching, often in fascicles, with
some intercalary segments inflated up to 15.0 um wide (cylindric), with walls thin or up
to 0.8 tum thick, with septa often constricted; acrophysalides dominating, narrowly fusi-
form to clavate to broadly clavate to ovoid to ellipsoid, up to 124 x 68 um, with walls
thin or up to 0.8 Um thick; vascular hyphae not observed; clamps not observed.
LAMELLA TRAMA: bilateral, with shallow to very shallow angle of divergence,
poorly rehydrating in some sections from the holotype, with central stratum 50 - 65 um
wide; filamentous, undifferentiated hyphae 1.8 - 9.2 um wide, branching, densely inter-
woven, with some intercalary segments slightly inflated; divergent, terminal, inflated
cells not observed; vascular hyphae not observed. SUBHYMENIUM: inflated ramose
to ramose, with depth of subhymenial tree 10+ tm under longest basidia/-oles and 25+*
uum under shortest basidioles, comprised of uninflated or partially inflated short hyphal
segments (up to about 16 tum long, but most shorter than 13 um long) in branching
structure, sometimes with ovoid to ellipsoid intercalary inflated cells (these larger than
most uninflated hyphal segments and concentrated adjacent to central stratum), rather
densely interwoven, with basidia arising from both ends and sides of hyphal segments,
with some segments parallel to the central stratum immediately below the bases of the
longest basidia, with about one to three cells between divergence from central stratum
and base of basidium. BASIDIA: 41 - 64 (-70) x (7.2-) 9.0 - 13.2 (-14.0) um,
dominantly 4-, but occasionally 2- or 1-sterigmate, many with yellowish brown to pale
yellowish brown contents in side view (orange-brown to red-brown in mass or in end
view) in both 3% KOH and 2-3% NH,OH, sometimes orange-brown to red-brown in
Melzer’s Reagent; clamps not observed. UNIVERSAL VEIL: On pileus (from
‘‘button’’): elements with vertical orientation except near base, there often with
periclinal orientation, frequently hyaline or pale yellow, occasionally orange-brown
(but then for only a few consecutive hyphal segments or even only for a portion of one
segment), less pigmented than adjacent pileipellis and pileus context; filamentous, un-
differentiated hyphae 3.2 - 15.0 (-26) um wide, branching, with walls thin or slightly
thickened, some with pale yellow walls, plentiful to dominating in the base of the wart,
otherwise scattered; inflated cells terminal, singly or in chains, broadly fusiform to
clavate to ovoid to ellipsoid to subglobose, with the roundest nearest top of wart,
dominating above base of wart, up to 112 x 64 um or larger, with walls usually at least
somewhat thickened (up to 0.8 tum thick); vascular hyphae 4.5 - 12.0 um wide, yellow-
brown, absent in most sections, relatively common in one, frequently branching; clamps
not observed. On stipe base: similar. STIPE CONTEXT: longitudinally acro-
physalidic, with cell walls hyaline to brown; filamentous, undifferentiated hyphae 1.5 -
10.5 um wide, branching, often in fascicles, plentiful, with walls thin or up to 1.0 ym
thick, with some intercalary cells having form and size of acrophysalides;
acrophysalides plentiful, up to 211 x 51 ym, with walls thin or up to 1.0 um thick; vas-
136
Figs. 3-6. Amanita westii. 3. Elements from base of universal veil from pileus of
‘“‘button’’ (Feibelman 1165). 4. Elements from upper portion of universal veil from
pileus of ‘‘button’’ (Feibelman 1165). 5. Elements of hymenium and subhymenium
and adjacent central stratum (Feibelman 1165). 6. Elements of partial veil (Lewis
4074). Dark bars indicate 20 um.
137
cular hyphae 4.0 - 21 tum wide, branching, yellow-brown, common. PARTIAL VEIL:
cells hyaline to yellow-walled to brown-walled; filamentous, undifferentiated hyphae
2.0 - 10.5 um wide, in tangled and interwoven fascicles, branching, with majority sub-
radially oriented, many collapsed or poorly rehydrating, plentiful to locally dominant,
with many terminal segments slightly inflated, with some intercalary segments slightly
inflated (cylindric), thin-walled or with walls slightly thickened; inflated cells fusiform
to narrowly fusiform to narrowly ellipsoid to subclavate, with narrower forms often
subrostrate to rostrate, up to 125 x 32 um, terminal, singly, with walls thin or up to 0.8
uum thick; vascular hyphae not observed.
BASIDIOSPORES: [100/4/3] (8.5-) 10.2 - 14.0 (-15.5) x (5.8-) 6.0 - 7.8 (-8.2) jum,
(L = 11.0 - 12.8 um; L’ = 12.1 um; W = 6.6 - 7.3 um; W’ = 7.0 um; Q = (1.42-) 1.50 -
1.99 (-2.18); Q = 1.67 - 1.79; Q’ = 1.73), hyaline, smooth, thin-walled, amyloid, ellip-
soid to elongate, often adaxially flattened, often slightly swollen at one end, often con-
stricted; apiculus sublateral, small, truncate-conic to cylindric; contents granular, in
holotype sometimes yellowish brown in dilute basic solution and turning reddish brown
in Melzer’s Reagent and then [per Bas (1969)] sometimes masking amyloid reaction;
color in deposit unknown.
Distribution and habitat: In Florida, subgregarious, under Quercus in dry, high
hammock (a term peculiar to Florida and indicating usually mesic, climax
vegetation——hardwood forest including Quercus, Magnolia, etc.—often slightly
elevated compared to surrounding terrain). In Mississippi, subgregarious in a mesic
forest of Pinus, Quercus, and Fagus near a large creek. In Texas, solitary in a xeric
forest largely of Quercus and Carya.
Collections examined: UNITED STATES: FLORIDA——Alachua Co. - east of
Gainesville, Newmann’s Lake, 7.vui.1938 E. West s.n. (holotype, FLAS F17466). MIS-
SISSIPPI——Perry Co. - De Soto National Forest, Cypress Creek Landing, 29.vi.1991
Toby Feibelman 1165 (F; L; RET). TEXAS——Newton Co. - Scrapping Valley Timber
and Wildlife Research Area, off State Hwy. 87 and Recreational Rd. 255, 1.viii.1987 D.
P. Lewis & J. Parigi [Lewis 4074] (F; RET).
DISCUSSION
The large, dark, pyramidal warts and reddish staining reaction of the context make
A. westii a striking species. It is unique in color and habit among the taxa of Amanita
described from the Western Hemisphere. The universal veil decorating the pileus often
takes on such a strikingly spiky form that, from a distance, one may mistake a specimen
of A. westii for a bolete belonging in the genus Strobilomyces B. Examination of the
universal veil on recently collected material (including ‘‘button’’ specimens) supports
the likelihood observed by Bas (1969; pers. comm.) that this North American entity has
as its closest known phyletic relative A. sculpta Corner & Bas (1962) described from
North Borneo and Singapore.
Gilbert (1941) suggested A. westii was synonymous with A. flavorubescens Atk.
The latter has a yellow to yellow-brown pileus with yellow, crumb-like warts and
belongs to Amanita section Validae (Fr.) Quél. Reddish bruising or staining reactions
138
in A, flavorubescens are usually restricted to context of the lower stipe and basal bulb.
In both 3% KOH and 2-3% NH,OH, the color of spore contents was not as strong as
observed by Bas (1969), but, instead, like the yellow of most refractive cell contents
seen in amanitas. The reddish-brown tint of spore contents observed in Melzer’s
Reagent was visible despite the strong amyloid reaction of the spore wall (observed at
1000x). However, the reddening of contents was never seen to occur so completely or
intensely as to mask the amyloid reaction as was noted by Bas. Bas noted scattered
brown vascular hyphae in the universal veil warts on the pileus; such hyphae were
found in only one of numerous sections of warts from recently collected material.
The stipe context of A. westii is curious in that many of the apparent acrophysalides
are seen to be intercalary cells with uninflated hyphal segments connected at both their
broad and narrow ends.
Tulloss has found (unpub. data) that yellow-walled, septate, filamentous, undif-
ferentiated hyphae are rather common in some taxa of Amanita section Lepidella. It is
possible that some authors have mistaken the pale yellow hyphae for vascular hyphae.
ACKNOWLEDGMENTS
We extend our gratitude to the following: Dr. Andrew S. Methven, Botany Depart-
ment, Eastern Illinois University, Charleston, for reviewing this paper; Ms. Toby
Feibelman, New Orleans, Louisiana, for her gift of the Mississippi material; Ms. Mary
A. King, Roosevelt, New Jersey, for assistance in preparing this paper for publication;
Mr. John Parigi, Beaumont, Texas, for assistance in collecting of the Texas material;
and Mr. David C. Tulloss, Berkeley, California, for preparing the habit illustrations.
LITERATURE CITED
Bas, C. 1969. Morphology and subdivision of Amanita and a monograph of its section
Lepidella. Persoonia 5(4): 285-579.
Comer, E. J. H. and C. Bas. 1962. The genus Amanita in Singapore and Malaya. Per-
soonia 2: 241-304.
Gilbert, E. J. 1941. Notules sur les amanites. (Libraire E. Le Francois, Paris). 23 pp. + 1
pl.
Jenkins, D. T. 1979. A study of Amanita types III. Taxa described by W. A. Murmill.
Mycotaxon 10: 175-200.
Munsell Color. 1976. Munsell Book of Color. Glossy Finish Collection. Baltimore.
unpaginated.
Murrill, W. A. 1944 [1945]. New Florida Fungi. Proc. Florida Acad. Sci. 7: 107-127.
Ridgway, R. 1912. Color standards and nomenclature. (self-published, Baltimore). 44
pp. + 53 pl.
Tulloss, R. E. 1993. Amanita pachysperma, Amanita subvirginiana, and Amanita vir-
giniana (taxonomy and distribution) with notes on description of the lamella trama
in Amanita. Mycotaxon 49: 449-475.
, C. L. Ovrebo and R. E. Halling. 1992. Studies on Amanita (Agaricales) from
Andean Colombia. Mem. New York Bot. Gard. 66: 1-46.
MY COTAXON
Volume L, pp. 139-150 January-March 1994
ORPINOMYCES INTERCALARIS, A NEW SPECIES
OF POLYCENTRIC ANAEROBIC RUMEN FUNGUS
FROM CATTLE
Y.W. HO
Department of Biology, Universiti Pertanian
Malaysia, 43400 UPM Serdang, Selangor, Malaysia.
N. ABDULLAH and S. JALALUDIN
Department of Animal Science, Universiti Pertanian
Malaysia, 43400, UPM Serdang, Selangor, Malaysia.
ABSTRACT
Orpinomyces intercalaris, a new species of polycentric
anaerobic fungus from the rumen of cattle is described and
illustrated. The zoospore germinates to produce
rhizomycelium which develops mostly intercalary sporangia.
The zoospore cyst is persistent and after germination
remains as an empty structure attached to the rhizomycelium.
It is very distinct during the early developmental stages of the
rhizomycelium. Sporangia are globose to subglobose.
Zoospores are polyflagellated.
The genus Orpinomyces was established by Barr et al. (1989)
with Orpinomyces bovis as the type species. The genus is
characterised by polycentric rhizomycelium with terminal or intercalary
branched sporangiophore complexes. Zoospores are polyflagellated
in contrast to Anaeromyces, the only other polycentric genus of
Neocallimasticales which has uni- to quadriflagellate zoospores
(Breton et a/, 1990.; Ho et al/., 1990; Ho et a/, 1993a). Only one
species (the type species) has been described and assigned to this
genus. The type species was originally named O. bovis Barr and Kudo
(Barr et al., 1989). However, slightly earlier, a polycentric fungus with
polyflagellate zoospores was described independently and assigned to
the genus Neocallimastix as N. joyonii Breton et al. (1989). The two
fungi remained separate taxa until Li et a/. (1991) studied the zoospore
ultrastructure and development of O. bovis and concluded that there
140
Fig. 1. A polyflagellated zoospore of O. intercalaris. Fig. 2. Zoospore
cyst with a long unbranched hypha (arrow). Figs. 3A-B. Zoospore cyst with a
short unbranched hypha (arrow). Bar = 20 um.
141
were not enough of distinctive differences between O. bovis and N.
joyonii to warrant separate genera. They also considered them to be
the same species. However, they concurred with Barr et al. (1989)
that a new genus, Orpinomyces, should be erected to accomodate this
new polycentric fungus. Since N. joyonii has priority over O. bovis, they
decided to combine the two names and renamed the fungus
Orpinomyces joyonii (Breton et al.) Li et al., comb. nov. (Li et a/.,
1991).
The fungus described in this paper was isolated from the
rumen contents of a cattle (Bos indicus) fed guinea grass (Panicum
maximum) ad libitum. The method of isolation was similar to that
described by Ho & Bauchop (1991). The media for culturing the fungal
isolate were glucose sloppy medium, cellulose sloppy medium and
liquid medium with rice straw as carbon source (Ho & Bauchop, 1991).
The isolate belongs to the genus Orpinomyces but is distinctly different
from O. joyonii.
Orpinomyces intercalaris Ho sp. nov. Figs. 1 - 10.
Rhizomycelium complexus, polycentricus et eucarpus. Hyphae
multiramosae, plerumque contrictae. Sporangia globosae vel subglo-
bosae subinde irregulares, intercalaria raro terminalibus. Zoosporae
globosae, subglobosae, vel forma variabiles, polyflagellatae. Cystae
zoosporae persistenti saepe ad rhizomycelium affixus. Specis obligate
anaerobia.
Rhizomycelium complex, polycentric and eucarpic. Hyphae
much-branched, often with constrictions. Sporangia globose or sub-
globose, occasionally irregular, intercalary rarely terminal. Zoospore
globose, subglobose or variable in shape, polyflagellate. Zoospore cyst
persistent, often attached to rhizomycelium. Obligate anaerobe.
From rumen contents of Kedah-Kelantan cattle (Bos indicus),
Universiti Pertanian Malaysia (UPM). Serdang, Malaysia, 25 March,
1993, Y.W. Ho. Type: Figs. 1-10. A culture, C70, is being maintained
at UPM.
The zoospores of O. intercalaris are mostly globose, about 8.5-
14 um in diameter, but occassionally subglobose or variable in shape
when in motion (Fig. 1). They are polyflagellated with about 12-20
flagella, 33-48 um long, posteriorly orientated. The flagella are shed
readily before encystment and germination. Shed flagella are
commonly seen next to the zoospore cyst and they usually possess a
142
Fig. 4. Initial development of an intercalary sporangium as a small
swelling in the hypha of a young rhizomycelium. A, swelling develops near
the zoospore cyst. B, swelling develops some distance from the zoospore
cyst. Figs. 5A-B. Initial development of intercalary sporangia as swellings in
the hyphae in older rhizomycelium. Bar = 20 um.
143
Fig. 6. Intercalary sporangium in a young rhizomycelium. A,
sporangium develops near the zoospore cyst. B-C, sporangium develops
some distance from the zoospore cyst. D, two sporangia develop some
distance from the zoospore cyst. Bar = 20 um.
144
Fig. 7. Intercalary sporangia in older rhizomycelium. A, sporangia
develop in one hypha. B, sporangium develops at a hyphal branch. C-D,
sporangia develop between a network of hyphae. E, sporangium with a
constricted neck (arrow). A-E, Bar = 20 um. F, a rhizomycelium with
intercalary sporangia and a persistent zoospore cyst (arrow) attached to it.
Bar = 100 um.
145
Fig. 8. Development of intercalary sporangium from one side of the
hypha. A, a young sessile sporangium develops as a swelling atthe side of
a hypha. B, mature sessile sporangia (arrows). C, young sporangium with a
bulbous basal structure. D, young sporangium with a cup-shaped basal
structure. E-F, mature sporangia with basal septum and empty cup-shaped
basal structure. G-H, irregularly-shaped sporangium with cup-shaped basal
structure. Bar = 20 um.
146
Fig. 9. Zoospore release. A, differentiated zoospores in a globose
sporangium. B, differentiated zoospores in an irregularly-shaped sporangium
with a cup-shaped basal structure. C, sporangium showing Zoospore release.
Note the flagella of zoospores emerging through the sporangial wall (arrow).
D-E, sporangium with rupture through which the zoospores are released.
Arrows show the released zoospores. F, empty sporangium (arrow) after
zoospore release. Bar = 20 um.
147
bead-like structure similar to that observed in Neocallimastix and
Piromyces species (Barr et.al.,1989; Ho et al., 1993b,c,d).
The zoospore germinates by producing a germ tube. The
contents of the zoospore are transferred to the germ tube which very
quickly expands and elongates to form a hypha. The hypha (3-9 um
diameter) can be unbranched for up to 250 um long before branching
to form a complex rhizomycelium (Fig. 2) or it can branch after a short
length (Figs. 3A & B). The empty zoospore cyst (5-9 x 6-20 um) is
persistent and remains as an empty structure attached to the
rhizomycelium (Figs. 2-4, 6). It is especially distinct during the early
developmental stages of the rhizomycelium.
Sporangia are mostly intercalary, very rarely terminal. They
usually develop as a small swelling (Figs. 4-5) in the hypha. The
swelling very rapidly expands and becomes separated from the hypha
following formation of two septa (Figs. 6C.7A). Sporangia may also
develop at points where hypha branch (Figs. 7B-F). The neck (the
part between the sporangium and hypha) is either wide (Figs. 7A-D) or
constricted (Fig. 7E). During the early developmental stages of the
rhizomycelium, sporangia can develop close to the zoospore cyst (Fig.
6A) or at some distance away (Figs. 6B-D). Occasionally, sporangia
develop from one side of the hypha instead of from the centre (Figs.
8A-D). Such sporangia can be sessile (Fig. 8B) or have a short
bulbous (Fig. 8C) or cup-shaped basal structure (Figs. 8D-F). Ina
mature sporangium, a basal septum is formed and the cup-shaped
structure appears empty (Figs. 8E-G). Sporangia with basal structures
are common in older cultures of 3-4 days in all the three media used.
Sporangia are mostly globose to subglobose (Figs. 6-8),
occasionally irregular, especially those with basal structures in older
cultures (Figs. 8G-H). Sporangium size is variable, ranging from 26-95
um in diameter. Most of the sporangia are concentrated at the centre
of the colony compose of rhizomycelium. There are fewer sporangia
at the edge of the colony.
Zoospore release is through rupture of the sporangial wall
(Figs. 9A-F). Collapse and dissolution of the sporangial wall take place
after zoospore release. Some of the sporangia, particularly the very
large ones, are apparently unable to release the zoospores. These
sporangia containing the zoospores become brown and moribund. A
similar phenomenon has been observed in some Neocallimastix
species (Barr et.al.,1989; Ho et al, 1993b). The thick wall in the larger
sporangia probably impede the normal release mechanism of the
sporangia (Barr, 1984).
Fig. 10. Rhizomycelium of O. intercalaris. A, hypha with a bead-like
appearance (arrow) with constrictions at close intervals. B, sausage-shaped
hyphae with constrictions at longer distance. C, tips of hyphae with fine to
moderately blunt ends. Bar = 20 um.
149
In glucose sloppy medium or cellulose sloppy medium, the
rhizomycelium grows extensively, forming discrete colonies of 2-4 mm
after 2-3 days. The rhizomycelium consists of much-branched, narrow
to wide hyphae ranging from 1.5-10.5 um in diameter. Some of the
larger hyphae have tightly constricted points at close intervals giving
the hyphae a bead-like appearance (Fig. 10A) or at longer intervals
resulting in sausage-shaped hyphae (Fig. 10B). The tips of the
hyphae are more or less straight with fine to moderately blunt ends
(Fig. 10C). Hyphae with tightly constricted points are very common in
polycentric anaerobic rumen fungi. They have been observed in O.
joyonii (= O. bovis, Barr et al., 1989) and Anaeromyces elegans Ho
(Ho et.al., 1993a) (= Ruminomyces elegans Ho, Ho et al., 1990). They
have also been observed in some monocentric anaerobic rumen fungi
such as Neocallimastix sp (Barr et. a/., 1989), N. variabilis (Ho et al.,
1993b) , Piromyces communis (Barr et.al., 1989) and P. minutus (Ho
et al., 1993c).
The presence of predominantly intercalary sporangia, which
develop between hyphae and not borne terminally on a
sporangiophore, is a distinct feature of O. intercalaris that distinguishes
it from O. joyonii. In O. joyonii, the sporangia are all terminal, formed
at the apices of sporangiophore complexes. The sporangiophores are
terminal or intercalary and they are irregularly branched. The
persistent empty zoospore cyst (after germination) which remains as a
distinct structure attached to the rhizomycelium is another
characteristic feature of O. intercalaris.
These distinct characteristics merit O. intercalaris to be
assigned to a new species.
ACKNOWLEDGEMENTS
We thank Dr. Ruth Kiew, Department of Biology, Universiti
Pertanian Malaysia, Serdang, Malaysia, for the Latin tanslation of the
diagnosis, and Dr. D.J.S. Barr, Centre for Land and Biological
Resources Research, Central Experimental Farm, Agriculture Canada,
Ottawa, Ont., Canada for reviewing the manuscript.
REFERENCES
Barr, D.J.S. (1984). The classification of Spizellomyces, Gaetnerio-
myces, Triparticalar and Kochiomyces (Spizellomycetales,
Chytridiomyces). Can. J. Bot. 62: 1171-1201.
150
Barr, D.J.S., Kudo, H., Jakober, K.D. and Cheng, K.-J. (1989).
Morphology and development of rumen fungi Neocallimastix
sp., Piromyces communis and Orpinomyces bovis gen. nov.,
sp. nov. Can. J. Bot. 67: 2815-2824.
Breton, A., Bernalier, A., Bonnemoy, F., Fonty, G., Gaillard, B. and
Gouet, P. (1989). Morphological and metabolic characteri-
zation of anew species of strictly anaerobic rumen fungus:
Neocallimastix joyonii. FEMS Microbiol. Letts. 58 : 309-314.
Breton, A., Bernalier, A., Dusser, M., Fonty, G., Gaillard-Martinie, B
and Guillot, J. (1990). Anaeromyces mucronatus nov. gen.,
nov. sp. A new strictly anaerobic rumen fungus with polycentric
thallus. FEMS Microbiol. Letts. 70: 177-182.
Ho, Y.W., Bauchop, T., Abdullah, N. and Jalaludin, S. (1990).
Ruminomyces elegans gen. et sp. nov., a polycentric anaerobic
rumen fungus from cattle. Mycotaxon 38 : 397-405.
Ho, Y.W. and Bauchop, T. (1991). Morphology of three polycentric
rumen fungi and description of a procedure for the induction of
zoosporogenesis and release of zoospores in cultures. J. Gen.
Microbiol. 137 : 213-217.
Ho, Y.W., Barr, D.J.S., Abdullah, N. and Jalaludin, S. (1993a).
Anaeromyces, an ealier name for Ruminomyces. Mycotaxon
47 : 283-284.
Ho, Y.W., Barr, D.J.S., Abdullah, N., Jalaludin, S. and Kudo, H.
(1993b). Neocallimastix variabilis, a new species of anaerobic
fungus from the rumen of cattle. Mycotaxon 46 : 241-258.
Ho, Y.W., Barr, D.J.S., Abdullah, N., Jalaludin, S. and Kudo, H.
(1993c). A new species of Piromyces from the rumen of deer
in Malaysia. Mycotaxon 47 : 285-293.
Ho, Y.W., Barr, D.J.S., Abdullah, N., Jalaludin, S. and Kudo, H.
(1993d). Piromyces spiralis, a new species of anaerobic
fungus from the rumen of goat. Mycotaxon 48 : 59-68.
Li, J., Heath, 1.B. and Cheng, K.-J. (1991). The development and
zoospore ultrastructure of polycentric chytridiomycete gut
fungus, Orpinomyces joyonii comb. nov. Can. J. Bot. 69 : 580-
589.
MY COTAXON
Volume L, pp.151-166 January-March 1994
THE ORIGINAL COLLECTIONS OF ARTURO NANNIZZI (1877-1961)
IN THE HERBARIUM UNIVERSITATIS SENENSIS (SIENA)
*CHIARUCCI A., °MARIOTTI M.G.
*Dipartimento di Biologia Ambientale, Universita di Siena,
Via P.A. Mattioli 4, I-53100 Siena (Italy)
°DIFCA, Facolta di Agraria, Universita di Milano,
Via Celoria 2, I-20133 Milano (Italy)
Abstract. the nature, size and actual state of conservation of the
Original mycological collection of Arturo Nannizzi (1877-1961),
eminent Siena mycologist of the Attilio Tassi (1820-1905)
school, are herewith illustrated. The complete autoptic material
of which Arturo Nannizzi described many new taxonomic units
is listed. Such material needs to be typified and this essay shall
represent a useful basis for those taxonomists interested in these
systematic groups.
Introduction
This paper deals with the description of the micromycetes
collections of the Herbarium Universitatis Senensis (SIENA). We believe a
catalogue of the material concerning the fungi described by Siena
mycologists useful for taxonomic study also due to the fact that the
specimens, often requested, have been returned with annotations but no
precise type designation. Hopefully specialists shall provide a typification of
this material and the collections shall be definitely and correctly arranged.
Care of the Siena collections has not been at the level of its importance in
the past and this has impeded the necessary support to systematic research of
value such as SUTTON's (1975; 1977). Increase of the collections’ value and
improvement of its management have recently been programmed (MARIOTTI
& CHIARUCCI, in press) and illustration of the original material to be
typified is the first step.
This first contribution examins those taxonomic units described by
Arturo Nannizzi (1877-1961). Pupil of Attilio Tassi (1820-1905), he became
152
a prominent mycologist developing an essential role in the study of many
mycetes mostly pathogenic to man (GIACOMINI, 1961; FERRI, 1987; RICCI,
1987). Only after his death and publications by GRIFFIN (1960),
SZATHMARY (1960), DAWSON & GENTLES (1961) and STOCKDALE (1961),
was his merit acknowledged: recognizing the dermatophyte as anamorphs of
ascomycetes belonging to the Gymnoascaceae family and hence placement in
the correct systematic position (NANNIZZI, 1926a). He described a new
genus of Hyphales (NANNIZZI, 1931) and many other new fungi species of
pathological interest to man (for a complete bibliography see GIACOMINI,
1961). He also worked profitably in the field of saprophyte fungi and plant
parassites of which he recognized and described many new species.
Arturo Nannizzi's collections in SIENA
The material studied by Siena mycologists of the Tassi school,
mostly fungi but also lichens (CHIARUCCI & Loppi, 1990) were originally
gathered by Flaminio Tassi in a collection denominated Mycotheca
Universalis. This was not only comparison material, holding also in fact
specimens originating from the Erbario Crittogamico Italiano, but also a
collection of the original specimens of taxonomic units described by Tassi.
The collection was later incremented by other mycologists with samples
from personal research, and with commercial and comparative collections as
the already cited Erbario Crittogamico Italiano, Mycotheca Italica by
Saccardo and the Mycotheca Germanica by Sidow. Presently the Mycotheca
Universalis holds 67 volumes of hard carton holders containing sheets on
which the specimen envelopes are pinned (MARIOTTI & CHIARUCCI, in
press).
Volumes 31-34, 41 and 43 contain the Nannizzi collection most of
which labeled with a reference note: 31: Funghi di Taranto (75 specimens);
32: Contributo Fl. Micol. della Bulgaria (48) e Ricerche sul Padule di
Fucecchio (48); 33: Mycologia Senese (80) e Batteriosi della Zinnia elegans
(1); 34: Funghi di Taranto, S. Nicolicchio (59); 41: Mycologia Senese (74);
43: Mycologia senese (45).
Autoptic material
Specimens belonging to the original Nannizzi collection, to be
typified, are listed here below. The list is based on the valid publicated
names (with eventual spelling corrections according to actual rules of
I.C.B.N.). The name, essential for the correct identification of the specimen,
is complete of bibliographic protologue quotation and description of the
collecting site. It is followed by the label wording of the specimen found in
P53
the Herbarium in inverted commas and, in square brackets, by the position
(of the volume and sheet) of the specimen itself in the "Mycotheca
Universalis" (SIENA).
For several reasons (author's handwriting, date, uniqueness of
almost all the specimens per species, etc.) we are convinced that the
specimens we found are the same ones Nannuzzi described; however, only
for a few did he specify that "[...] the material examined is preserved in the
Mycotheca of the Botanical Institute of Siena [...]". In addition for some
taxonomic units, already validly described in NANNIZZI (1926b) and
NANNIZZI (1927) the author repeats after the diagnosis including an
autographed iconography (NANNIZZI, 1928a) which could be used for
typification according to the art. 9.3 of the I.C.B.N.. The various
possibilities of typification hence should be studied.
We believe that the micromycetes should have a detailed
designation of the type, clearly showing what corrisponds to the protologue
on the autoptic material and including eventual microscope slides and
iconographies as the type. The example of Clathrospora patriniae Nannizzi
(see further ahead) evidences this belief. Many are the cases in which what
appears a holotypus at first glance, is really a number of elements partly
used for the protologue and partly foreign to it. Although difficult and not
usual, we think a choice (lectotypification) of that or those sole elements
indispensable and sufficient. This choice should not only be stated in
revision papers but also clearly indicated on the collection material.
The original material on whose basis NANNIZZI (1925b)
redescribed the genera Heterobotrys Sacc. illegitimately modifying the name
to Morularia Nannizzi, has unfortunately been lost. According to MEECKER
(1975) Heterobotrys paradoxa Sacc. (= Morularia saccardiana Nannizzi) is
the anamorph of Seuratia millardetii (Raciborski) Meecker.
List of the described taxonomic units:
ASCOMYCOTINA
DOTHIDEALES
Mycosphaerellaceae
Sphaerella agostinii Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 437-
438 (1927).
Hab. in foliis vaginisque exsiccatis Crithmi maritimi in parva insula S.
Nicolicchio prope Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Sphaerella crithmi sp. n./ Foglie e guaine secche di Crithmum/ maritimum.
Isolotto di S. Nicolicchio/ (Taranto) Sett. 1927" (SIENA, [34: 18]).
Chianicci A Manatti MG 4
154
Note: as known, the genus Sphaerella (Fr.) Rabenh. non Sommerf. (Algae)
& nomen gen. illegitt. and is substituted by Mycosphaerella Johanson (1884).
Sphaerella sassafras Ell. & Ev. f. major Nannizzi, Atti R. Acc. Fisiocr.
Siena, ser. X, 4: 89 (1929).
Hab. in foliis emortuis Sassafras officinalis in Horto botanico Senensi,
Italiae. Oct. 1928 (NANNIZZI, 1929).
"Sphaerella Sassafras Ell. et Ev./ forma major Nannizzi/ Foglie di Sassafras
officinale/ Orto bot. Siena/ Ottobre 1928 A. Nannizzi" (SIENA [33: 32]).
Note: see previous species.
Pleosporaceae
Clathrospora patriniae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 960
(1928).
Hab. in foliis petiolisve putrescentibus Patriniae sibiricae in Sibiria australi,
M. Karakol, 4 jun. 1923 (NANNIZZI, 1928c).
"Clathrospora patriniae Fol. putrescenti di Patrinia sibirica" (SIENA [41:
35]).
Note: Clathrospora is nomen gen. rejic. in favour of Pleospora,
nevertheless SHOEMAKER & BABCOCK (1992) examining the original
material of the species described by Nannizzi, form the new combination
Platysporoides patriniae (Nannizzi) Shoemaker & Babcock. In a recent and
accurate paper, these authors identify four different taxonomic units of the
Original material including two petioles and a small leaf fragment. The
characters of two of the taxonomic units were used for the original
description; in fact Nannuzzi probably exchanged ascospores of Pleospora
comata Auerswald & Niessl for mature stages of Clathrospora spores. With
valid motivations do SHOEMAKER & BABCOCK (1992) maintain the specific
name limiting it to the taxonomic unit with applanate ascospores.
Leptosphaeria altaica Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 961
(1928).
Hab. in vaginis exsiccatis foliorum Polygoni Bistortae in Sibiria australi
prope Cemal secus flum. Katum, M. Altai, 20 jun. 1923 (NANNIZZI, 1928c).
"Leptosphaeria_altaica sp. n./ Guaine secche di Polygonum/ Bistortae"
(SIENA [34: 9]).
Note: Leptosphaeria is nomen gen. cons.
Pleospora herbarum (Pers.: Fr.) Rabenh. f. rosae-banksiae Nannizzi, Riv.
Pat. Veg., 18 (9-10): 190 (1928).
Habitat in foliis vivis Rosae Banksiae in Horto botanico Senensi, Italiae; oct.
1928 (NANNIZZI, 1928b).
155
"Pleospora herbarum Rabh./ f. rosae Banksiae Nannizzi/ Sulle foglie di Rosa
Banksia/ Orto bot. di Siena/ 17 ottobre 1928 A. Nannizzi" (SIENA [33:
31)).
ERYSIPHALES
Erysiphaceae
Erysiphe polygoni DC. ex Mérat f. robiniae-hispidae Nannizzi, Riv. Pat.
Veg., 14 (9-10): 173-174 (1924).
Hab. in foliis vivis vel languentibus Robiniae hispidae in horto botanico
Senensi, mense octobri 1924. Status conidicus orbiculari sistit (NANNIZZI,
1924).
Note: no specimen found in SIENA. Previously, NANNIZZI (1923) had
described the conidic stage of the taxonomic unit denominating it, in a way
now illegitimately, Oidium orbiculare (= O. monosporum Passerini). No
sample was found in SIENA, not even under this binomio.
SPHERIALES
Trichosphaeriaceae
Eriosphaeria dumetorum Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
438 (1927).
Hab. in ramulis siccis Rosmarini officinalis in dumetis prope Chiatona,
Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Eriosphaeria littoralis sp. n./ Rametti di Rosmarinus officinalis/ 7 sett. 27.
Chiatona, lungo il mare" (SIENA [31: 6]).
Xylariaceae
Anthostomella maritima Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
437 (1927).
Hab. in tunicis siccis bulborum Urgineae maritimae in parva insula S.
Nicolicchio prope Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Anthostomella maritima sp. n./ Tuniche bulbi di/ Urginea maritimae/ S.
Nicolicchio/ Taranto/ Sept. 1927" (SIENA [31: 1]).
DEUTEROMYCOTINA
COELOMYCETES
Ascochytella vaginarum Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
439 (1927).
Hab. in vaginis exaridis foliorum Pini halepensi, Chiatona, Tarentum. Sept.
1927 (NANNIZZI, 1927, 1928a).
"Diplodina vaginarum/ sp. n./ Guaine foliari di Pinus halepensis/ Chiatona.
Sett. 1927" (SIENA [31: 5]).
156
Note: the genus Aschochytella Tassi has been erroneously typified in the
past as A. deformis (Karst.) Died. (CLEMENTS & SHEAR, 1931), species not
included among the thirteen species originally described by TAssI (1902).
This has already been evidenced by SUTTON (1977).
Camarosporium gnidii Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 441
(1927).
Hab. in ramis emortuis Daphne Gnidii in dumetis prope Chiatona, tarentum.
Sept. 1927 (NANNIZZI, 1927, 1928a).
"Camarosporium Gnidii sp. n./ Rametti secchi di Daphne Gnidium/ nella
macchia di Chiatona. Sett. 1927" (SIENA [41: 35]).
Camarosporium obtusum Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 1:
506 (1926).
Hab. in scapis, calycibus bracteisve siccis Statice Limonii var. serotinae in
parva insula S. Nicolicchio pr. Tarentum, Italia austr. Aug. 1926
(NANNIZZI, 1926b, 1928a).
"Camarosporium obtusum/ Scapi, calici e brattee di Statice/ Limonium L.
var. serotina Rchbg./ Taranto aug. 1926" (SIENA [34: 2]).
Camarosporium psammae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
441 (1927).
Hab. in culmis siccis Psammae arenariae in arenosis prope Storranum,
Punta S. Vito, Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Camarosporium Psammae sp. n./ Culmi secchi di Psamma arenaria/
Chiatona (Taranto) Sett. 1927" (SIENA [31: 1]).
Coniothyrium garryae Nannizzi, Arch. Bot. Sist. Fitogeogr. Genet., 8: 298
(1932);
Hab. in ramulis emortuis Garryae Faydeanae in horto botanico Senensi cult.
Nov. 1931 (NANNIZZI, 1932).
"Coniothyrium Garriae n. sp./ Rametti secchi di Garrya Faydena/ orto bot.
Siena Nov. 1931" (SIENA [41: 36]).
Note: species not cited by BIGA et al. (1959).
Coniothyrium manihotis Nannizzi, Arch. Bot. Sist. Fitogeogr. Genet., 8:
298 (1932).
Hab. in ramis emortuis Manihotis palmatae in horto botanico Senensi. Nov.
1931 (NANNIZZI, 1932).
"Coniothyrium Manihotis/ n. sp./ rami morti di Manihot palmata/ Ort. bot.
Siena Nov. 1931" (SIENA [41: 35]).
Note: species not cited by BIGA et al. (1959).
157
Coryneum feijoae Nannizzi, Arch. Bot. Sist. Fitogeogr. Genet., 8: 300
(1932).
Hab. in foliis vivis Feijoae Sellowianae in horto botanico Senensi cult. Febr.
1932 (NANNIZZI, 1932).
"Coryneum Feijoae n.sp./ foglie vive di Feijoa Sellowiana/ Orto bot. Siena.
Febbr. 1932" (SIENA [41: 35]).
Note: SUTTON (1975) supposes that the species be referred to the genus
Seimatosporium Corda, but precises that not having received the original
material from SIENA for accurate examination he doubts the identity of the
species described by Nannizzi.
Cryptosporium lunatum Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
442 (1927).
Hab. in ramulis emortuis Pinus halepensi in sylvis dumetisque prope
Chiatona, Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Crysptosporium lunatum sp. n/ (mescolato ad un ascomicete rametti secchi
di Pinus halepensis/ Chiatona (Taranto) Sett. 1927" (SIENA [41: 36]).
Note: SUTTON (1980) distributes the various species of Cryptosporium
among different genera, but C. /unatum is not cited.
Didymochaeta atropae Nannizzi, Atti R. Acc. Fisiocr. Siena, sez. Agr., 5:
37-38 (1938).
Hab. in caulibus emortuis Atropae Belladonnae in montibus Orlovdol,
Bulgariae (NANNIZZI, 1938a).
"Didymochaeta Atropae/ in caulibus emortuis Atropae Belladonnae/ 1936"
(SIENA [53: 10]).
Note: in SIENA (Mycotheca Universalis) vol. 32 (where Bulgaria specimens
are preserved), at pag. 26 there is a label indicating: "10/ Didymochaeta
atropae Nannizzi n. sp.", but the specimen is missing. A specimen can be
found at page 10 of vol. 53 (Mycologia Senese), where the material of
another research is preserved.
Diplodia dorycnea Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 440
(1927).
Hab. in ramulis emortuis Dorycnii hirsuti in dumetis prope Chiatona,
Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Diplodia dorycnea n. sp./ Rametti secchi di Dorycnium hirsutum/ Chiatona
pr. Taranto Sept. 1927" (SIENA [31: 4]).
Diplodia porlieriae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 5: 387
(1930).
158
Hab. in ramis emortuis Porlieriae hygrometricae in Horto Botanico Senensi;
jul. 1930 (NANNIZZI, 1930).
"Diplodia porlieriae Nann./ rametti morti di Porlieria hygrometrica/ Ort.
Bot. Siena. Aprile 1930/ (Nannizzi)" (SIENA [43: 14]).
Diplodia smilacella Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 440
(1927).
Hab. in foliis putrescentibus Smilacis asperae in dumetis prope Chiatona,
Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Diplodia _smilacella sp. n./ Fol. putresc. di Smilx aspera/ Chiatona
(Taranto) Sett. 1927" (SIENA [31: 5]).
Diplodina odontitis Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 439
(1927).
Hab. in caulibus siccis Odontitis luteae in nemoribus prope Chiatona,
Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Diplodinia Odontitidis n. sp./ con associata Didymosphaeria_ brunneola
Niessl./ Macchia di Chiatona su cauli secchi di/ Odontites lutea. Sett. 1927"
(SIENA [31:5]).
Hendersonia cyperi-aegyptiaci Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X,
2: 440 (1927).
Hab. in calamis foliisque siccis Cyperi aegyptiaci in dunis maritimis prope
Chiatona, Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Hendersonia arenicola sp. n./ sulle foglie di Cyperus aegyptiacus/ Chiatona
(Taranto)/ Sept. 1927" (SIENA [34: 9])
"Hendersonia arenicola sp. n./ Scapi di Cyperus aegyptiacus/ sulle arene
marittime pr. Chiatona/ (Taranto) Sett. 1927" (SIENA [31: 6)).
Note: Hendersonia Sacc. 1884 non Berk 1841 is nomen gen. rejic. pro
Stagonospora, but some species have been included in Hendersonula. A
reliable identity cannot be attributed to the species described by Nannizzi
without examination of the original material.
Hendersonia pteleae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 5: 387
(1930).
Hab. in ramis emortuis Pteleae trifoliatae in Horto Botanico Senensi; oct.
1929 (NANNIZZI, 1930).
"Hendersonia pteleae Nann./ Rami morti di Ptelea trifoliata/ Orto bot. di
Siena. Ott. 1929/ (Nannizzi)" (SIENA [43: 13]).
Note: see previous species.
Tov
Macrophoma leptopoda Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
439 (1927).
Hab. in ramis siccis Nicotianae glaucae, S. Nicola prope Punta Rondinella,
Tarentum. Aug. 1927 (NANNIZZI, 1927, 1928a).
“Macrophoma Nicotianae sp. n./ Cauli morti di Nicotiana glauca/ S. Nicola,
sui muri, Ag. 1927/ (Taranto) (SIENA [34: 10]).
Note: Macrophoma is nomen gen. rejic. pro Sphaeriopsis Sacc. nomen cons.
(SUTTON, 1980).
Pestalotia ventricosa Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 442
(1927).
Hab. in ramulis siccis Lonicera implexae in dumetis prope Chiatona,
Tarentum. Sept. 1927 (NANNIZZI, 1927, 1928a).
"Pestalozzia ventricosa n. sp./ Rametti secchi di Lonicera implexa/ associata
a Pleospora herbarum/ Chiatona (Taranto) Sett. 1927" (SIENA [31: 12]).
Note: there are two schools: one holds a wide concept of the genus (GUBA,
1961) the other subdivides the genus in Pestalotia, Pestalotiopsis and
Truncatella (STEYAERT, 1949, 1953, 1955; SUTTON, 1969). Only the
examination of the original material could allow a convincing identification
of the species described by Nannizzi.
Phaeoseptoria rubiae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 441
(1927).
Hab. in foliis vivis Rubiae peregrinae in dumetis prope Chiatona, Tarentum.
Sept. 1927 (NANNIZZI, 1927, 1928a).
"Phaeoseptoria Rubiae Nann./ Sulle foglie di Rubia peregrina/ Chiatona
(Taranto) Sett. 1927" (SIENA [34: 11]).
Phoma debeauxi Roum. f. scapicola Nannizzi, Atti R. Acc. Fisiocr. Siena,
ser. X, 2: 438 (1927).
Hab. in scapis emortuis Statice Limonii var. serotinae prope S. Annam,
Punta S. Vito, Tarentum. Aug. 1927 (NANNIZZI, 1927, 1928a)
"Phoma_ debeauxi Roum. forma/ Scapicola n.f./ Scapi secchi di Statice
Limonium var. se-/ rotina. S. Vito alla Casa dei Bagnanti. Ag. 1927"
(SIENA [32: 13]).
Phoma fimbriata Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 438
(1927).
Hab. in caulibus emortuis Inulae crithmoidi in parva insula S. Nicolicchio
prope Tarentum Sept. 1927 (NANNIZZI, 1927, 1928a)
"Phoma fimbriata sp. n./ Cauli morti di Inula crithmoides/ S. Nicolicchio.
Sett. 1927/ (Taranto)" (SIENA [34: 12]).
160
Phoma herbarum Westd. var. loti-cretici Nannizzi, Atti R. Acc. Fisiocr.
Siena, ser. X, 1: 505 (1927).
Hab. in caulibus emortuis Loci cretici in parva insula "S.Nicolicchio" pr.
Tarentum, Italia austr. Aug. 1926 (NANNIZZI, 1926b, 1928a).
"Phoma herbarum/ var. Loti-cretici Nannizzi 1926/ Rametti secchi di Lotus
creticus/ Chiatona (Taranto) 7 sett. 1927" (SIENA [31: 15)).
Phoma schini-molli Nannizzi, Atti R. Acc. Fisiocr. Siena, Ser. X, 2: 439
(1927).
Hab. in ramulis siccis Schini Molli cult. in horto "Villa Peripato",
Tarentum. Sept. 1927. Legit Doct. A. Agostini (NANNIZZI, 1927, 1928a).
"Phoma Schini-Molli sp. n./ Rametti di Schinus Molle/ "Villa Peripato",
Taranto. Sept. 1927" (SIENA [31: 15]).
Phomopsis phellodendri Nannizzi, Arch. Bot. Sist. Fitogegor. Genet., 8:
297 (1932).
Hab. in ramis emortuis Phellodendri amurensi in horto botanico Senensi
cult. Nov. 1931 (NANNIZZI, 1932).
"Phomopsis Phellodendri n. sp./ Rami secchi di Phellodendron amurense/
Orto bot. Siena Nov. 1931" (SIENA [32: 13]).
Phyllosticta aberiae Nannizzi, La Vedetta Agricola 2, 14 (1912).
Hab. in foliis vivis Aberiae caffrae in horto botanico Senensi culta, mense
novembri 1909 (NANNIZZI, 1912).
"Phyllosticta Aberiae Nannizzi/ Sulle foglie di Aberia caffra/ Orto botanico
di Siena/ Aprile 1912 A. Nannizzi" (SIENA [33: 29]).
Phyllosticta altaica Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 961
(1928).
Hab. in foliis vivis Veronicae incanae in Sibiria australi secus flum. Matun,
M. Altai, 24 jun. 1923 (NANNIZZI, 1928c).
"Septoria altaica n. sp./ Foglie vive di Veronica incana/ Siberia (M. Altai)
(SIENA [34: 17]).
Note: Only one leaf of Veronica, very small and with very rare pustules,
makes up the specimen. The corrispondence of the specimen to Phyllosticta
(Phoma s.\.) or Septoria has not been controlled to avoid its destruction;
however, due to the considerable differences between two genera, this
identification would be useful.
Phyllosticta helwingiae Nannizzi 1930, Atti R. Acc. Fisiocr. Siena, ser. X,
4: 91 (1929).
161
Hab. in foliis vivis Helwingiae rusciflorae in Horto botanico Senensi,
Italiae. Sept. 1927 et 1928 (NANNIZZI, 1929).
"Phyllosticta Helwingiae Nannizzi/ Sulle foglie di Helwingia ruscifolia/ Orto
botanico di Siena/ Settembre 1928 A. Nannizzi" (SIENA [33: 30]).
Note: two specimens.
Phyllosticta ibotae Nannizzi Atti R. Acc. Fisiocr. Siena, ser. X, 2: 438
(1927).
Hab. in foliis vivis Ligustri Ibotae cult. in horto prope Tarentum. Aug. 1927
(NANNIZZI, 1927, 1928a).
"Phyllosticta Ibotae n. sp./ Agosto 1927 Taranto/ Ligustrum Ibota/ Siebl. et
Zucc./ (var. Regelianum e Obovatum)" (SIENA [31: 14]).
Phyllostictella draconis Nannizzi, Arch. Bot. Sist. Fitogegr. Genet., 8: 297
(1932).
Hab. in foliis vivis Dracaena Draconis in horto botanico Senensi cult. Oct.
1930 (NANNIZZI, 1932).
"Phyllostictella Dracaenae/ n. sp./ Sulle foglie vive di Dracaena Draco/ Orto
bot. Siena Ott. 1930" (SIENA [34: 11]).
Note: according to SUTTON (1977) the genus Phyllostictella Tassi (1901)
should probably be considered synonymous of Microsphaeropsis Moehn
(1917).
Rhabdospora rubiae Pat. var. amerospora Nannizzi, Atti R. Acc. Fisiocr.
Siena, ser. X, 2: 441 (1927).
Hab. in foliis siccis Rubiae peregrinae in dumetis prope Chiatona, Tarentum.
Sept. 1927 (NANNIZZI, 1927, 1928a).
“Rhabdospora rubiae Pat./ var. amerospora n. var./ Foglie secche di Rubia
peregrina/ Macchia di Chiatona. Sett. 1927/ (Taranto)" (SIENA [34: 17]).
Robillarda aquatica Nannizzi, Boll. Pesca Piscic. Idrobiol., 14: 807 (1938).
Hab. in foliis putrescentibus natantibus Lemnae minoris in padule
Fucecchio, Florentiae; 15 mart. 1932 (NANNIZZI, 1938b).
"Robillarda aquatica n. sp./ Foglie putrescenti di Lemna minor/ Padule di
Fucecchio/ 15-II-32" (SIENA [32: 55]).
Note: according to SUTTON (1980) Robillarda Cast. (1845) should be
rejected in favour of Pestalotiopsis Stey (1949); while for the ICBN
(GREUTER, 1988), Robillarda Sacc. is nomen gen. conserv.
Septoria asteris-alpinis Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 961
(1928).
162
Hab. in foliis siccis Asteris alpini in Sibiria australi prope Cemal, secus
flum. Katun, M. Altai, 9 jun. 1923 (NANNIZZI, 1928c).
"Septoria Asteris-alpini sp. n./ Foglie secche di Aster alpinus" (SIENA [34:
17]).
Septoria iridis-japonicae Nannizzi, Arch. Bot. Sist. Fitogeogr. Genet., 8:
299 (1932).
Hab. in foliis vivis Iridis japonicae in horto botanico Senensi cult. Apr.
1931 et 1932 (NANNIZZI, 1932).
"Septoria Iridis-japonicae/ n. sp./ Sulle foglie vive di Iris japonica/ Ort. bot.
Siena/ apr. 1932" (SIENA [34: 18]).
Septoria moesiaca Nannizzi, Atti R. Acc. Fisiocr. Siena, sez. Agr., 5: 38
(1938).
Hab. in foliis Atropae Belladonnae in montibus Buzludgia, Bulgariae
(NANNIZZI, 1938a).
Note: missing specimen. At pag. 21 of vol. 32 of the Mycotheca
Universalis, an only sheet reports "Septoria moesiaca Nannizzi n. sp./ 21".
The specimen was probably lent and never returned.
Septoria oligocarpa Nannizzi, Riv. Pat. Veg., 15: 3 (1925)
Hab. in foliis vivis Cobeae scandentis in horto botanico Senensi culta, mense
decembri 1924 (NANNIZZI, 192S5a).
Note: no specimen found.
Septoria pachypleuri Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 962
(1928).
Hab. in foliis siccis vel languescentibus Pachypleuri alpini in Sibiria australi,
M. Karakol, 17 jun. 1923 (NANNIZZI, 1928c).
"Septoria Pachypleuri et/ Sphaerella Pachypleuri Fuck. in/ Heuglin Reisen
Nordpol., Vol. III, et/ Oud. Contr. Fl. Myc. Now. Seml. p. 6,/t. II. f. 2 -
Sacc. Syll. IX. p. 624/ Foglie secche di Pachypleurum alpinum" (SIENA
[34: 18]).
Stagonospora schoeni Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2: 440
(1927).
Hab. in calamis emortuis Schoeni nigricans in arvis arenosis pr. "Salina
Piccola", Tarentum Aug. 1927 (Nannizzi, 1927, 1928a).
"Stagonospora Schoeni sp. n./ Calami morti di Schorenus nigricans/ campi
arenosi alla Punta S. Vito, Taranto/ (mescolato con altri micromiceti) Ag.
1927 (SIENA [31: 25}).
163
Torula altaica Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 3: 962 (1928).
Hab. in petiolis putrescentibus foliorum Saxifragae crassifoliae in Sibiria
australi, M. Karakol, 17 jun. 1923 (NANNIZZI, 1928c)
"Torula altaica sp. n./ Picciuoli fogliari di Saxifraga/ crassifolia" (SIENA
[34: 21]).
HYPHOMYCETES
Macrosporium phoenicis Nannizzi, Atti R. Acc. Fisiocr. Siena, ser. X, 2:
439 (1927).
Hab. in foliis emortuis Phoenicis reclinatae cult in horto "Villa Peripato",
Tarentum. Aug. 1927 (NANNIZZI, 1927, 1928a).
"Macrosporium Phoenicis n. sp./ Foglie secche di Phoenix reclinata/ Villa
Peripato, Taranto 1927" (SIENA [31: 10]).
Note: according to HUGHES (1958) Macrosporium Fr. (1832) non Fr. (1825)
Should be synonymous of Alternaria Nees (1816/1817). This was confirmed
by ELLIs (1971).
Cladosporium anonae Nannizzi, Atti R. Acc. Fisiocr. Siena, ser X, 4: 91
(1929).
Hab. in foliis vivis Anonae sp. in Horto botanico Senensi, Italiae; Aug.
1928 (NANNIZZI, 1929).
"Cladosporium Anonae Nannizzi/ Sulle foglie di Anona sp./ Orto Botanico
di Siena/ Agosto 1928 A. Nannizzi" (SIENA [33: 27]).
Acknowledgements
The authors are grateful to Prof. Aurora Corte Montemartini for
comments and suggestions and to Mrs. Nancy Jane van Laarhoven for help
during the preparation of the manuscript.
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MY COTAXON
Volume L, pp. 167-174 January-March 1994
LAMBERTELLINIA SCUTULOIDES (SCLEROTINIACEAE),
A NEW GENUS AND SPECIES
FOR A DISCOMYCETE PREVIOUSLY CONFUSED WITH
HYMENOSCYPHUS CAUDATUS
RICHARD P. KORF & PAVEL LIZON
Plant Pathology Herbarium, Cornell University
Ithaca, NY 14853-4203, USA
Abstract
A phaeosporous member of the Sclerotiniaceae, a new
genus and species, Lambertellinia scutuloides Korf &
Lizon, with a substratal stroma and an Idriella
anamorph, is described.
KEYWORDS: Sclerotiniaceae, Lambertellinia, L. scutu-
loides, Leotiaceae, Hymenoscyphus caudatus, Idriella.
During a survey of the genus Hymenoscyphus an unidentified
collection of a helotioid Discomycete from Japan was studied. It has
some similarities in spore shape and size to Hymenoscyphus caudatus
(P. Karst.) Dennis. Another collection, which we treat as conspecific
with the Japanese collection, was published under that name by
Kimbrough & Atkinson (1972). The Japanese collection is
distinguished from the members of Hymenoscyphus, as understood by
us (Lizon, 1992), in having light brown spores and a different
construction of the excipular structures, and in producing a substratal
stroma. We propose it here as a new species and a new genus in the
family Sclerotiniaceae.
168
Lambertellinia scutuloides Korf & Lizon, gen. & sp. n.
= (Helotium berberidis Sydow, Mycoth. March. no. 1576, 1887 (nom.
nud.)
Misappl.: Hymenoscyphus caudatus (P. Karst.) Dennis sensu
Kimbrough & Atkinson, Amer. J. Bot. 59:165-171, 1972;
sensu Dennis, Mycol. Pap. 62:82, 1956, pro parte.
Apothecia stipitata, lutea, solitaria, stromatis
substratalibus. Discus plano-concavus, luteus vel
aurantiacus. Receptaculum cupulare, stramineum vel
aureum, ad 3.5 mm diam. Stipes concolor, basi
brunnescens, 1-4 mm longus et 0.2-1 mm crassus.
Excipulum ectale exterius ex textura porrecta bene evoluta
formatum (5-20 um latum), hyphis stramineis vel aureis,
0.5 - 1.5 Um diam., excipulum ectale interius ex textura
porrecta vel oblita formatum (ca. 15 um latum) hyphis
flavidis, stratum intermedium (35-55 um latum) ex hyphis
hyalinis et refractivis, 3-6.5 um diam., muris gelatinosis
formatum. Excipulum medullare ex textura porrecta vel
intricata formatum (ca. 20 um latum), hyphis parallelis vel
intertextis, 1.5 um diam. Asci cylindrico-clavati, octospovi,
(82-) 84.5-100 (-116) x (5.6-) 6.6.-7.5 (-8.3) um, poro in
Melzero caerulescente, non ex uncis orientes. Paraphyses
filiformes, 1.5 Um diam. et ascos non transeuntes.
Ascosporae oblique biseriatae, clavatae (“scutuloideae’) vel
inaequilaterales, laeves, primum hyalinae et non-septatae,
demum intra ascum pigmentum manifestantes, brunneae
vel hyalinae, saepe 1-septatae, (13.8-) 15.4-16.9 (-18.5) x
(2.3-) 3.1-3.5 (-3.8) um. Holotypus: CUP-JA 2252.
Apothecia yellow, long-stipitate, arising from a black substratal
stroma, scattered. Receptacle deep cupulate, with a sharp margin,
pale yellow to golden yellow (to brownish), slightly furrowed towards
the stipe, up to 3 mm diam. Disc concave to plane, darker yellow
than the outer surface to orange-yellow. Stipe cylindric, pale yellow to
golden-yellow (to brownish), sometimes brownish towards the base,
mainly longer than the diameter of the receptacle, 1-4 x 0.2-1 mm,
usually with a black ring of stromatal tissue in the flesh of the base.
169
Ectal excipulum of 2-3 layers of textura porrecta. Outermost
covering layer (5.0-20 um wide) of parallel, straw- or golden-yellow
pigmented hyphae 0.5-1.5 um diam., outer ectal excipulum (35-55 um
wide) of parallel hyphae 3.0-6.5 um, hyaline with glassy gelatinized
walls, inner ectal excipular cells light yellow with more visible walls
(layer about 15 um wide). Medullary excipulum of textura porrecta
to textura intricata, forming a layer about 20 um wide on the flanks,
Lambertellinia scutuloides, an apothecial section of holotype
specimen at CUP: ME - medullary excipulum, EC -
medullary excipulum, ie - inner layer of ectal excipulum,
ge — gelatinous layer of ectal excipulum, ce — covering layer
of ectal excipulum.
170
composed of parallel or slightly interwoven hyaline hyphae 1.5 um
diam. Asci cylindric-clavate, conical at the apex, (82—-) 84.5-100
(-116) x (5.6-) 6.6-7.5 (-8.3) um, pore weakly blue in Melzer’s reagent,
not arising from croziers. Paraphyses filiform, 1.5 um diam., not
extending beyond the asci. Ascospores biseriate or irregularly
uniseriate, clavate to inequilateral, “scutuloid’! hyaline and
non-septate when young, light brown or hyaline and occasionally 1-
septate when mature and when discharged, with 1-4 (-5) guttules
when inasci, but usually eguttulate when discharged, smooth, (13.8-)
15.4— 16.9 (-18.5) x (2.3-) 3.1-3.5 (-3.8) um. Stroma substratal, well
developed, rind-like, covering nearly the whole surface of petioles,
evident as a superficial blackening of the substrate, in transverse
section as a blackened zone surrounding the epidermal cells, in facial
view as blackish intercellular lines.
HOLOTYPE: Japan: Honshu, Tsugawa, Ogunimachi, Yamagata Pref.,
500-800 m, On Aesculus turbinata, 28. VIII.1961, D. Shimizu (CUP-JA
2252).
ETYMOLOGY: Lambertellinia refers to the relationship to
Lambertella, and scutuloides refers to the shape of ascospores, similar
in shape to those of Hymenoscyphus caudatus and H. scutulus.
SUBSTRATA: Petioles of Acer, Aesculus, and Bevberis.
SPECIMENS EXAMINED: JAPAN: Honshu, Tsugawa, Ogunimachi,
Yamagata Pref., 500 - 800 m, On Aesculus turbinata, 28. VIII.1961, D.
Shimizu (holotype of Lambertellinia scutuloides, CUP-JA 2252).
[GERMANY], Auf Berberis vulgaris, Hippodrom bei Berlin, VIII.1887,
P. Sydow (Sydow, Mycoth. March. no. 1576, 1887: FH, NY, “syntypes’
of Helotium berberidis Syd.).
A culture derived from a collection of a discomycete on leaves of
Acer rubrum, identified by Kimbrough & Atkinson (1972) as
Hymenoscyphus caudatus, produced an Idriella anamorph. It is
clearly different from Hymenoscyphus caudatus. The original
specimen is not available but according to the description ascospores
' The term “scutuloid” was introduced by Baral (Baral & Krieglsteiner, 1985) for
clavate spores in the genus Hymenoscyphus with the apex rounded and often
slightly hooked to one side, and the base pointed.
a |
developed 1-2 septa before germination, and a brown coloration.
Both these characters exclude it from Hymenoscyphus in the current
sense. Based on the description and excellent illustrations of the
characters of the excipulum, spores and asci we believe their
collection belongs to Lambertellinia scutuloides. Kimbrough and
Atkinson's misinterpretation of H. caudatus was followed by
Hennebert & Bellemére (1979), Kendrick & DiCosmo (1979), von Arx
(1981), Rappaz (1992), and Rodrigues & Samuels (1992).
Sydow has issued in his Mycotheca Marchica Helotium berberidis
Syd. (no. 1576, 1887), but no description was provided on the
exsiccati label. This species has apparently never been described. The
collection has apothecia dark brown (nearly blackish) when dry,
brownish when rehydrated, outer surface yellowish, stipe yellow, thin,
about half as long as the diameter of the receptacle. Apothecia arise
from stromatized veins and petioles of leaves. The ectal excipulum is
of textura porrecta, including a layer of hyphae with gelatinized walls.
Asci are clavate, conical at the apex, pore weakly blue in Melzer’s
reagent. Paraphyses are filiform, not exceeding the ascus tips, 2,2 um
diam. Ascospores are biseriate, cylindric clavate, “scutuloid’, deep
brown to hyaline, non-septate, 16.2 x 4.5, 16.8 x 4.8, 17.1 x 4.5, 17.4 x
42, 18.0 x 4.8, 19.8 x 3.6 um. [all data derived from dry material].
These characters clearly show that this collection (‘species’) is
conspecific with Lambertellinia scutuloides as described here, since
the difference in apothecial color when dry may represent only the
conditions under which they were dried, and the age of the
specimens.
Dennis (1956) examined specimens of Helotium berberidis in Sydow’s
Mycotheca Marchica and noted that it “is H. caudatum but is
apparently a nomen nudum.” He overlooked the brown coloration of
spores. We have searched for a description of this species but we
have found only a note on the publication of this exsiccatum (Bot.
Zeitung 46:204, 1888). A “syntype” specimen at FH has several well
developed apothecia, while the “syntype” at NY consists mostly of
broken leaves bearing no apothecia.
172
TAXONOMIC POSITION OF LAMBERTELLINIA
Though there are some genera of the Leotiaceae which produce
brown ascospores, or tardily brown ascospores, none of these
(Phaeohelotium, Bulgaria, Bulgariella, Sorokina, Polydiscidium,
Velutarina, Pestalopezia, Trochila, Catinella, among others) has an
excipular structure at all recalling that of Lambertellinia, and all lack
a substratal stroma. The stroma is the clearest indication that we are
dealing here with a member of the Sclerotiniaceae. Substratal
stromata occur in many genera of the Sclerotiniaceae, and normally
are an indication that infection of the substrate has occurred while it
was still living. Among the genera with brown or brownish
ascospores and a substratal stroma are currently only Lambertella and
Phaeociboria (if that is considered distinct, synonymized by Dumont,
1971). The new genus differs markedly from species of Lambertella in
the long-celled structure of the ectal excipulum and, in particular, in
the production of an Idriella anamorph (no species of Lambertella is
known to produce a conidial anamorph).
There are species in the Hymenoscyphus subserotinus “group,” mostly
tropical, which may prove to be more closely related to
Lambertellinia scutuloides than to other members of Hymenoscyphus.
Like Lambertellinia, they display a more or less strongly gelatinized or
glassy, long-celled excipular structure, and ascospores that are
frequently scutuloid (though not regularly turning brown), and there is
some evidence of a substratal (but limited) stroma. None have been
shown to produce a conidial anamorph. They appear to represent an
interface between Leotiaceae (Hymenoscyphoideae) and
Sclerotiniaceae. One species in this group, Lanzia serotina (Pers. : Fr.)
Korf & Zhuang, has recently been shown to produce an extensive
stroma in culture but no conidial anamorph (Zhuang, 1993).
Equally puzzling are two species of Crocicreas (or of Cyathicula if
that is considered distinct from Crocicreas as suggested by Baral &
Krieglsteiner, 1985) collected by the senior author in Macaronesia.
Pure cultures of these two collections produce a substratal stroma in
some, but not all, culture media. It seems probable that other
members of cyathiculoid fungi may also represent an interface
between Sclerotiniaceae and Leotiaceae. For example, Cyathicula
hyalina Beaton has been transferred to Poculum (or, more correctly,
to Rutstroemia) on the basis of the presence of a well-developed
173
substratal stroma. Unlike typical Rutstroemia spp., however, this has
spores brown at maturity, according to Carpenter (1981). The senior
author has unpublished data on European collections of a common
Rutstroemia species that also develops brown pigmentation on spore
germination.
In addition, many (mainly tropical) species placed in such genera as
Hymenoscyphus and Cyathicula are simply insufficiently investigated
to determine whether a substratal stroma is produced. For most, no
cultures exist to prove the point, or to demonstrate the presence of a
conidial anamorph.
Acknowledgments
We wish to thank Dr. Linda Kohn (University of Toronto) for
reviewing this paper and Dr. William J. Dress (Cornell University) for
the revision of the Latin diagnosis. This research was supported by
Anna E. Jenkins bequest in a form of a postdoctoral fellowship to the
junior author.
Literature cited
Arx, J. A. von, 1981: Notes on Microdochium and Idriella. Sydowia
34:30-38.
Baral, H. O. & Krieglsteiner, G. J., 1985: Bausteine zu einer
Ascomyceten-Flora der Bundesrepublik Deutschland: In
Siiddeutschland gefundene inoperculate Discomyzeten. Beih. Z.
Mykol. 6:1-160.
Dennis, R. W. G., 1956: A revision of the British Helotiaceae in the
Herbarium of the Royal Botanic Gardens, Kew, with notes on
related European species. Mycol. Pap. 62:1-216.
Carpenter, S. E., 1981: Monograph of Crocicreas (Ascomycetes,
Helotiales, Leotiaceae). Mem. New York Bot. Gard. 33:1-290.
Hennebert, G. L. & Bellemére, A., 1979: Les formes conidiennes des
Discomycétes. Essai taxonomique. Rev. Mycol. (Paris)
43:259-315.
174
Kendrick, B. & DiCosmo, F., 1979: Teleomorph-anamorph
connections in Ascomycetes. In: Kendrick, B. (ed.), The whole
fungus 1:283-359, Ottawa.
Kimbrough, J. W. & Atkinson, M., 1972: Cultural features and
imperfect state of Hymenoscyphus caudatus. Amer. J. Bot.
59:165-171.
Lizon, P., 1992: The genus Hymenoscyphus (Helotiales) in Slovakia,
Czechoslovakia. Mycotaxon 45:1-59.
Persoon, C. H., 1801: Synopsis methodica fungorum. Pars 1. p. [II}/- XXX,
(1}+-240, pars 2, o. [241}-706, [I-II], Gottingae [Gottingen].
Rappaz, F., 1992: Phomatospora berkeleyi, P. arenaria and their
Sporothrix anamorphs. Mycotaxon 45:323—330.
Rodriguez, K. F. & Samuels, G. J., 1992: Idriella species endophytic in
palms. Mycotaxon 43:271-276.
Zhuang, W.-y., 1993: Lanzia serotina in culture. Acta Mycol. Sinica 12:
85-86.
MY COTAXON
Volume L, pp. 175-189 January-March 1994
AGARICALES OF BAJA CALIFORNIA - MEXICO
M. Candusso! - A. Gennari2 - N. Ayala?
1 Via Grandi 28 I-21047-Saronno
2 Via Anconetana 35/a I-52100 Arezzo
3 Laboratorio de Micologia-Universidad Autonoma de Baja California-Mexico
Abstract
A preliminary agaric checklist covering all the species collected is presented.
Four of them: Clitocybe pinophila, Omphalina rosella, Lactarius
argillaceifolius var. megacarpus and L. alnicola are fully described and
discussed.
Key words: Fungi, Basidiomycotina, Agaricales, Russulales, Clitocybe,
Omphalina, Lactarius, Clitocybe pinophila, Omphalina rosella, Lactarius
alnicola, L. argillaceifolius var. megacarpus.
Introduction
A large number of fresh fruiting bodies of Agaricales were collected and
identified during a brief stay in Baja California - Ensenada - Mexico. Of the
23 species listed 4, considered to be especially interesting, were described in
detail. The state of Baja California has an amazing floral diversity due to a
wide range of habitats, going from sand dunes to oak woods up to the pine
forests at 1600 m above sea level. The climate is rather similar to that typical
of the Mediterrancan area, with very hot summers and relatively cool winters.
Prior to our stay the rainfall in the single month of January had been
exceptional, this circumstances affected badly our expedition to Sierra Juarez.
Materials and Methods
Microscopic examination, measurements and drawings were made using a
Bausch & Lomb and an Olympus BH2 microscopes, equipped with
planapochromatic lens and drawing tubes. Freehand sections were mounted in
2% KOH and stained in Melzer's reagent and/or congo red. The material
studied is deposited partly in the herbarium of UABC-Ensenada-Mexico and
partly in the herbarium of the first author.
Taxonomy
Bolbitius vitellinus (Pers.: Fr.)Fr. var. fragilis (L.: Fr.), Ep. Syst. Myc.: 254, 1838.
176
PLAYA HERMOSA - ENSENADA-B.C.: 12-02-93 leg. et det.: M.C.
Habitat : on sandy soil between the sea and a lagoon. Exs. N° 93021204.
Clitocybe fragrans (With.: Fr.) Kummer, Der Fihrer in die Pilzk.: 121, 1871.
PLAYA HERMOSA - ENSENADA-B.C.: 12-02-93 leg. et det.: M.C.
Habitat : on sandy soil between the sea and a lagoon. Exs. N° 93021205.
Clitocybe pinophila (Peck) Saccardo, Syll. Fung. V: 183, 1887.
Agaricus (Clitocybe) pinophilus Peck, Rep. New York State Mus. 31: 32,
1879.
ASERRADERO-RANCHO LAS BOTELLAS SIERRA JUAREZ: 17-02-93 leg.: C.
Sigitenza Lopez, det.: MLC.
Habitat : on needles of Pinus jeffreyi, granitic soil, elev. approx. 1520 mt. Exs.
N° 93021702.
Original description
Agaricus (Clitocybe) Pinophilus Pk.
Pileus thin, convex, umbilicate or centrally depressed, glabrous, moist, pale
tan-color, paler or alutaceous when dry; lamellae moderately close,
subarcuate, adnate or slightly decurrent, whitish; stem equal, stuffed or
hollow; glabrous or subpruinose, colored like the pileus; spores nearly
elliptical, .0002-.00025 long; odor and taste resembling that of fresh meal.
Plant 1 -2' high, pileus about 1' broad, stem 1" -2" thick.
Ground under pine trees. Albany and Ticonderoga. July and August.
Description
Pileus 1,5-3 cm broad, plane, plano-convex, depressed at centre, then
infundibuliform; surface glabrous, looking waxy, hygrophanous; margin thin,
slightly projecting; date-brown, brownish-yellow, pale brown, buff, ochre-
brown, honey color, pallescent on drying, margin paler.
Lamellae from adnate to broadly adnate or slightly decurrent, rather crowded,
anastomosed and intervenose mostly near the lamellulae, L/l = 1/3-5; cream-
ochre color, sub-concolor to pileus, hygrophanous.
Stipe 1,5-4 x 0,2-0,5 cm, cylindrical, sometimes compressed, often flexuose,
spindle-shaped, slightly swollen towards base, sometimes with white rhizoids
at the base of the stem; surface smooth, minutely fibrillose; pale brown, pale
fawn, sub-concolor with pileus, paler specially at apex.
Context pale brownish, hygrophanous, sub-concolor with surface, smell
spontaneously often subnitrous or weak, taste none.
40 um
Stipitipellis,
trama, st
pileipellis, tr
Clitocybe pinophila: pi
ba
spores.
basidia, sp =
178
Habitat on needles of Pinus jeffreyi.
Microscopic features
Spores (4,5)-5-6,4 x 2,8-3,5 yum, ellipsoid, oval, smooth, hyaline, negative in
Melzer's reagent. Basidia subcylindric, clavate, stout, often bubble-shaped
towards base (up to 3,5-5 ym), with clamp connections, (18)-20-23 x 4,5-6
pm, 4-spored, (rarely 2-spored).
Pileipellis a cutis with cylindrical cells, irregular, 25-75 x 5-10 pm, with
several clamp connections. Pigment intracellular, yellow-brownish.
Stipitipellis a cutis of cylindrical hyphae, with rounded terminal elements 75-
140 x 5-10 pm.
Trama of the gills made of parallel to subparallel hyphae, 70-120 x 5-12 ym,
with several clamps.
Clamps present in all tissues.
Observations: In Peck's description there is no mention of the intervenose or
anastomosed lamellae we observed in our specimens. Clitocybe subbulbipes
shows much the same macroscopical features, but has different habitat. The
latter is very close to C. americana but differs in its color and the presence of a
finely encrusting pigment in the pileipellis. C. californiensis is distinct by the
absence of intervenose lamellae and habitat (Quercus agrifolia). All these
species grow under hardwood trees while the former fruits under conifers
(Pinus jeffreyi).
It is also close to C. acerba Bigelow, but the latter is readily distinguished by
its whitish color and smaller spores. The smell as noted by us was not
farinaceous but weakly nitrous.
A good color reproduction of C. pinophila can be seen in the recently
published "Mushrooms of North America” by R. Philips under the name of C.
subbulbipes.
Clitocybe trullaeformis (Fr.: Fr.) P. Karst., Ryssl. Finl. Skand. Halféns
Hattsvamp. 1: 72, 1879.
LAS CHICHIHUAS-KM. 80 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93 leg.: G. Moreno-M.C., det. M.C.
Habitat : on soil with Rhus integrifolia, Quercus agrifolia, elev. approx. 420
mt. Exs. N° 93021002.
Coprinus lagopides P. Karst., Ryssl. Finl. Skand. Halféns Hattsvamp. 1: 535,
1879.
LA PINTA-SAN QUINTIN: 13-02-93 leg. et det.: M.C.
Habitat : on sandy soil near the sea. Exs. N° 93021302
179
Entoloma sericeum (Bull. ex) Quél. var. sericeum, Mém. Soc. Emul.
Montbéliard, sér. II, 5: 119, 1872.
LAS CHICHIHUAS-KM. 80 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93 leg.: M.C., det.: A. Galbusera
Habitat : on grass near Rhus integrifolia, elev. approx. 420 mt. Exs. N°
93021006
Entoloma lividoalbum (Kiihn. & Romagn.) Kubicka, Ceska Mykol. 29: 27,
1975.
CANADA VERDE-KM. 106 CARRETERA TECATE MEXICALI: 06-02-93
leg. et det.: M.C.
Habitat : Quercus agrifolia, elev. approx. 1000 mt. Exs. N° 93020602.
Laccaria bicolor (R. Mre.) Orton, Trans. Br. mycol. Soc. 43: 177, 1960
CANON DE STA.ROSA-KM.78 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93 leg. et det.: M.C.
Habitat : Quercus agrifolia near the Sta. Rosa's river, elev. approx. 420 mt.
Exs. N° 3021005.
Lactarius alnicola A.H. Smith var. alnicola, Brittonia 12: 319, 1960.
IGNACIO ZARAGOZA-KM..65 CARRETERA LIBRE-ENSENADA-TECATE:
06-02-93 leg. et det. A.G.
Habitat: Quercus agrifolia, elev. approx. 350 mt. Exs. N° 93020609.
Original description
Pileus 8-15 cm latus, convexo-depressus, glutinosus, luteus, ad marginem
pubescens; sapor forte acris; latex albus; lamellae tactu luteae. Specimen
typicum legit A.H. Smith (N° 59648), 5 Aug. 1958, Warm Lake, Idaho, U.S.A.
(MICH).
Description
Pileus 5-12(18) cm, at first convex, broadly depressed, margin inrolled and
slightly velvety, in age often nearly plane, surface slightly viscid, zonate when
young, ochraceous yellow, paler at the margin, like L. zonarius.
Lamellae adnate-decurrent, arcuate, close to crowded, narrow, cream whitish,
many lamellulae present.
Stipe 3-7 x 2-3 cm, cylindrical, broadened at apex, somewhat smaller at the
base, stuffed then hollow, pubescent, whitish to cream yellowish, with rare
yellow scrobiculation.
Context white, yellow when cut, taste at first mild becoming somewhat acrid
only after some minutes. Latex white, copious, turning citron yellow instantly.
ae
sp
—_
ch
Lactarius argillaceifolius var. megacarpus: sp = spores,
ch = cheilocystidia
a | ca
AG.
Lactarius alnicola var. alnicola: pi = pileipellis, sp = spores,
ch = cheilocystidia, ca = caulocystidia
pl
181
KOH yellow on pileus cuticle, citron yellow on flesh.
Habitat: under Quercus agrifolia.
Microscopic features
Spores 6,5-9,5(10) x 6-9 ym, ellipsoid to subglobose, with a partial reticulum,
prominences relatively high.
Basidia 4-spored, clavate, 40-50 x 9-13 pm.
Cheilocystidia (30)-50-90 x 3-8 jm, clavate, narrow, with acuminate apex.
Caulocystidia in KOH yellow.
Pileipellis an ixocutis of 3-6 ym wide, encrusted hyphae.
Observations: The european L. zonarius is very close, but is readily
distinguished by the whitish latex. Other species of Lactarius which resemble
our species are: L. payettensis, L. yazoonensis, L. olympianus, L. psammicola
Jm. glaber, L. alnicola var. pitkinensis and L. alnicola var. pungens.
Lactarius argillaceifolius Hesler & Smith var. megacarpus Hesler & Smith
North Am. Sp. of Lactarius: 369, 1979.
KM.15 AL PARQUE NACIONAL COSTITUCION DEV.KM. 55 -
ENSENADA-S.FELIPE: 17-02-93 leg.: A.G.-M.C.-C. Sigiienza-M. Lizarraga,
det. A.G.-M.C.
Habitat: Quercus agrifolia, Quercus dummii, Rhus integrifolia, elev. approx.
1100 mt. Exs. N° 93021701
Original description
Pileus 14-27 cm latus, viscidus, azonatus, luteolus vel ad marginem pallido-
avellaneus. Latex albus, constans, branchias colore brunnaceotingens.
Lamellae adnatae, paene subdistantes, luteolae. Stipes 16-20 cm longus, 40-
5O mm crassus, leviter viscidus, basi albus, apice sordide luteolus. Sporae
7.5- 10.5 x 6.5-8 uu. Specimen typicum in Herbarium University of Tennessee,
Thiers 18556 (TENN 35424), legit Prope Santa Barbara, California, 28 Jan
1967.
Description
Pileus 12-22 cm, convex, plano-convex, regular to very irregular, with deep
central depression. Pileal covering smooth, slightly and finely velvety when
dry, margin hardly viscid when moist, strongly inrolled, extremely wavy,
irregular, indented, pubescent and strongly striate. Often with soil particles
overall. Grayish violet, lilac lead-gray, with buff tinges, disc with lighter
182
ochraceus color. Margin silvery gray. Reminiscent of Russula cyanoxantha.
Lamellae adnate, slightly decurrent, at times forked at stipe attachment,
crowded. L/] = 1/5-7, of medium thickness, narrow. Argillaceous cream, dark
cream, staining brownish, vinaceous gray when touched. Latex very acrid after
about 10 seconds, cream to ivory cream.
Stipe 8-12 x 3-6-(8) cm, cylindrical, slightly ventricose, very irregular,
canaliculate, compressed, tapering at base, hollow, stuffed. Surface smooth,
finely pubescent, hardly viscid when wet, at times scrobiculate, whitish
ochraceous-cream, staining brownish yellow.
Context tough in outer layer of stipe, pithy-cottony inside, white, cream,
sometimes almost pink, subconcolor under the cuticle. Taste slightly acrid
after a short while, odor slightly Pentalomide-like (cimicic).
Habitat under Californian oaks, gregarious to caespitose.
Microscopic features
Spores 8-9(10) x 6,5-7,5 jm, ellipsoid, ornamentation of fine lines not
forming a complete reticulum.
Basidia 4-spored.
Cheilocystidia fusoid with acuminate apex, irregular, with one or more
constriction toward the apex.
Pileipellis an ixotrichodermium formed by a tangle of hyphae 2.5-4.5 ym
wide, with rounded and/or capitulate end-cells, some dissepiments and
branchings with yellowish intracellular pigment and pieces of blackish brown
extracellular pigment.
Observations: The pileipellis structure, undoubtedly trichodermial according
to our observation, does not fit with the original description.
Macrolepiota rachodes (Vittadini) Singer var. bohemica (Wichansky) Bellu
& Lanzoni, Beitr. Kenn. Pilze Mittel. III: 191, 1987.
RANCHO LOS JACARANDAS-CANON LAS ANIMAS: 07-02-93 leg.: N.
Ayala-M. Lizarraga, det.: N.A.-M.C.
Habitat : under Sambucus mexicanus, elev. approx. 200 mt. Exs. N°
93020708.
Micromphale inodorum (Pat.) Dennis, Kew Bull. 15, 1: 90, 1961.
LAS CHICHIHUAS-KM. 80 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93 leg.: A.G., det.: A.G.-M.C.
Habitat : Heteromeles arbutifolia, elev. approx. 420 mt. Exs. N° 93021004.
183
Nothopanus cf. lignatilis (Pers.: Fr.) Bon
RANCHO LOS JACARANDAS-CANON LAS ANIMAS: 07-02-93 leg. et
det.: M.C.
Habitat : on Opuntia litoralis, elev. approx. 200 mt. Exs. N° 93020704.
Omphalina rosella (Lge.) Moser, Helmut Gam's KI]. Krypt. Mitt. II: 58, 1953.
Basionymum ( = Omphalia rosella Lge., Dansk Bot. Arkiv 6(5): 14).
RANCHO LOS JACARANDAS-CANON LAS ANIMAS: 07-02-93
Habitat : on grasslands near Q. agrifolia, elev. 200 mt. Exs. N° 93020701.
LAS CHICHIHUAS-KM. 80 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93
Habitat : on grassland, elev. approx. 420 mt. Exs. N° 93021018.
KM.71-SAN QUINTIN-STA.INES: 13-02-93
Habitat : grass on granitic soil, elev. approx. 30 mt. Exs. N° 93021301, leg. et
det. M.C.
Original description
Small. Cap 0.7-1.5cm., membranaceous, at first umbilicate with incurved
edge, then infundibuliform, of a pale and somewhat dingy pink color (without
striation), centre slightly dirt-brownish. The edge is somewhat irregularly
crenato-fimbriate and slightly rugoso-plicate. When dry the color is paler and
duller. Gills deeply decurrent rather narrow, somewhat furcate, rather distant,
pale pink. Stem short (1-2 cm. x 1% mm.), at first pinkish, then whitish,
glabrous, somewhat wavy, not particularly tough and not hollow.
Spores cylindrical-ellipsoid, obliquely pedicellate, 712-9 x 4-412 yu. Basidia 4-
spored. Cystidia scattered, short hairshaped about 4 yu thick (the base, which
is imbedded in the tissue of the gill, about 6 y).
Fig. specim. (D.A. 292): "Slukefter” near Odense, in a park, gregarious on old
lawn, (light soil), Oct. 1921 (and 23).
Pileus diametro 0.7-1.5 cm., membranaceus, umbilicatus (margine incurvato)
dein infundibuliformis, laevigatus, roseolus (pars centralis sordide subfulva),
margine (in adultis) crenato-fimbriato vel rugoso-plicato, pallescens; lamellis
longe decurrentibus, angustis subfurcatis, subdistantibus, pallide roseolis.
Stipes brevis (1-2 cm. x 1% mm.), roseolus (in adultis albus), glabratus.
Sporae et cystidia ut supra. -
Description
Pileus 0,5-1,5-(2) cm broad, convex to concave, usually deeply umbilicate,
infundibuliform, with slightly to distinctly involute margin when young, then
straight or reflexed and fringed in old specimens; surface slightly velvety, dry,
184
color very variable, first dirty white or pale ochre, then whitish cream,
yellowish cream, pinkish white, pinkish ochre, with centre brownish when
mature.
Lamellae deeply decurrent, arcuate, anastomosed, rather distant, 2- 3-furcate,
L/l = 1/1-3, white, cream, pinkish, then ochre with tender pinkish tinge, with
heteromorphous and sometimes pink or brown edge.
Stipe 2-3 x 0,1-0,2 cm, cylindrical, solid, smooth, dry, sometimes broadened
at base, subconcolor with pileus.
Habitat terrestrial in groups in grasslands, often on rather poor, sandy soil;
near Quercus agrifolia and Skinus mollis.
Microscopic features
Spores (6,5)-7-9-(9,5) x 4,5-5,5 ym, subamygdaliform, subpyriform, ellipsoid,
with evident hilar apiculus, smooth. Basidia clavate, subcylindrical, 30-35 x 7-
9 pm, 4-spored, sterigmata 4-5 x 1,5-2 ym wide, with clamp connections at
base. Cheilocystidia making the edge heteromorphous, conspicuous, clavate,
mucronate, apically forked (obtuse), cylindrical, irregular, hyaline.
Pileipellis a cutis with subcylindrical, subellipsoid elements, with rounded,
sometimes free terminal cells, of variable diameter and rather versiform 30-
250 x 5-15 pm.
Stipitipellis a cutis with elements 50-75 x 5-7,5 jm, sometimes with
subcylindrical, free elements, with rounded terminal cells. Clamp connections
present at every septum.
Trama of the gills with mostly filamentous and regular hyphae, rather dense,
with subparallel arrangement, formed mainly by subcylindrical, subellipsoid
elements 25-40 x 5-9 ym.
Clamps present in all tissues.
Observations: O. rosella resembles O. ericetorum (= O. umbellifera) but
differs mainly by the absence of clamp connections and the presence of
cheilocystidia. In addition it differs from O. subclavata (Peck) Murrill among
other characters by the total absence of yellow-olive tinges. Another species
that is very close, but has white colors and habitat in coniferous woods is O.
californiensis. O. jalapensis has different spore-size (5 x 3,5 jum).
Murrill published several species of Omphalia in 1916 (O. Mac Murphyi, O.
Bakeri, O. tepeitensis, O. niveicolor ecc.), but we were unable to find our
species among them because the descriptions are either very incomplete or too
short.
Omphalotus olivascens Bigelow, Miller & Thiers var. indigo Moreno, Esteve-
Raventos, Péder & Ayala (Mycotaxon in print).
CANON DE STA.ROSA-KM.78 CARRETERA LIBRE-ENSENADA-
Omphalina rosella: tr = trama, ch = cheilocystidia, ba = basidia,
Sp = spores, st = stipitipellis, pi = pileipellis
185
186
TIJUANA: 10-02-93 leg.: M.C., det.: G. Moreno-M.C.
Habitat : Quercus agrifolia, elev. approx. 420 mt. Exs. N° 93021003.
Observations: For a long time this species was confused with O. olearius or
O. illudens. The abundance of specimens of O. olivascens in all stages of
development (with pileus diameter up to 35 cm) collected in Baja California,
along with our previous experience of both Mediterranean species, enabled us
to confirm the validity of the species proposed by Bigelow, Miller & Thiers in
Mycotaxon (1976).
Pluteus nanus (Pers.: Fr.) Kummer, Der Fiihrer in die Pilzk.: 98, 1871.
PLAYA HERMOSA-ENSENADA:; 12-02-93 leg. et det.: M.C.
Habitat : Haplopapus sp., Pluchea sericea on marsches with grass. Exs. N°
93021201.
Psilocybe coprophila (Bull.: Fr.) Kummer, Der Fihrer in die Pilzk.: 71, 1871.
LAS CHICHIHUAS-KM. 80 CARRETERA LIBRE-ENSENADA-TIJUANA:
10-02-93 leg. et det.: M.C.
Habitat : on dung, elev. approx. 420 mt. Exs. N° 93021001.
Tricholoma columbetta (Fr.: Fr.) Kummer, Der Fiihrer in die Pilzk.: 131,
1871.
KM.15 AL PARQUE NACIONAL COSTITUCION DEVIACION KM.55-
ENSENADA-S.FELIPE: 17-02-93 leg.: G. Giana, det.: A.G.
Habitat : Quercus agrifolia, elev. approx. 1100 mt. Exs. N° 93021703.
Volvariella gloiocephala (DC.: Fr.) Boekh. & Enderle, Beitr. Kenntn. Pilze
Mitteleur. 2: 78, 1986.
RANCHO LOS JACARANDAS-CANON LAS ANIMAS: 07-02-93 leg. et
det.: A.G.-M.C.
Habitat : on grasslands, elev. approx. 200 mt. Exs. N° 93020703.
Acknowledgements
We gratefully acknowledge the Universidad Autonoma de Baja California for
the assistance and for providing additional financial support. During this visit
we had the assistance of Prof. G. Moreno and Dr. C. Ochoa in identifying
some of the described species.
We would like to express our appreciation to members of the Universidad de
Alcala de Henares - Madrid : Dr. Miriam Blanco, Dr. C. Illana and Dr. R.
Galan; to the members of Universita di Trieste : F. Bersan, Dr. G. Giana, Dr.
M. Sarasini and Amer Montecchi for collecting fungi.
187
Thanks also to Biol. C. Sigiienza Lopez for the determinations of host plants
and for supplying valuable field information and personal observations.
A special note of gratitude is due to M. Lizarraga and Biol. J. Delgadillo who
have provided us valuable collections and field data.
All of these people were also wonderful partners during various trips to several
localities in Baja California.
We wish to thank Prof. Dr. M. Moser for the review of this manuscript.
References
Arora, David 1979: Mushrooms Demystified. Berkeley, Ca.
Ayala, Nahara & G. Guzman 1984: Los hongos de la Peninsula de Baja
California. I. Las especies conocidas. Boletin de la Sociedad Mexicana de
Micologia 19: 73-91.
Bigelow, Howard E. 1958: New species and varieties of Clitocybe from
Michigan. Mycologia 50: 37-51.
Bigelow, Howard E. 1970: Omphalina in North America. Mycologia 62: I-
32:
Bigelow, Howard E. 1977: New taxa of Clitocybe. Mycotaxon 6: 181-185.
Bigelow, Howard E. 1982: North American Species of Clitocybe. Part I.
Beihefte zur Nova Hedwigia 72.
Bigelow, Howard E. 1985: North American Species of Clitocybe. Part II.
Beihefte zur Nova Hedwigia 81.
Bigelow, H. & A.H. Smith 1962: Clitocybe species from the Western United
States. Mycologia 54: 498-515.
Bigelow, H.-O.K. Miller Jr.-H.D. Thiers 1976: A new species of
Omphalotus. Mycotaxon 3: 363-372.
Boekhout, Teun 1990: Volvariella in Flora Agaricina Neerlandica 2: 56.
Guzman, Gastén 1973: Some distributional relationships between Mexican
and United States mycofloras. Mycologia 65: 1319-1330.
Guzman, Gastén 1980: Identificacién de los hongos. Limusa, D.F., Mexico.
Hesler, L.R. & A.H. Smith 1979: North American Species of Lactarius. The
University of Michigan.
Hesler, L.R. & A.H. Smith 1960: Studies on Lactarius-II. The North
American species of sections Scrobiculus, Crocei, Theiogali and Vellus.
Brittonia 12, N° 4: 306-350.
Kauffman, C.H. 1918: The Agaricaceae of Michigan. 2 Vols. Johnson
Reprint.
Laferriére J.E. & R.L. Gilbertson 1992: Fungi of Nabogame, Chihuahua,
Mexico. Mycotaxon 44: 73-87.
Lamoure, D. 1974-75: Agaricales de la zone alpine. Genre Omphalina.
Travaux scientifiques du Parc National de la Vanoise V-VI.
Lange, Jakob E. 1930: Studies in the Agarics of Denmark. Omphalia.
188
Pleurotus. Clitocybe. Dansk Botanisk Arkiv 6 (5): 1-19.
Lange, Jakob E. 1935: Flora Agaricina Danica. Copenhagen.
Moser, Meinhard 1978: Kleine Krypt. Flora. Band II, Teil b 2. Gustav
Fischer.
Mueller, Gregory M. 1992: Systematics of Laccaria (Agaricales) in the
Continental United States and Canada, with Discussions on Extralimital Taxa
and Descriptions of Extant Types. Fieldiana (1435) 30: 1-158.
Murrill, William A. 1913: The Agaricaceae of the Pacific Coast-IV. New
species od Clitocybe and Melanoleuca. Mycologia 5: 206-223.
Murrill, W.A. 1912: The Agaricaceae of the Pacific Coast-Il. Mycologia 4:
231-262.
Murrill, W.A. 1912: The Agaricaceae of the Pacific Coast-III. Mycologia 4:
294-308.
Orr, Robert T. & Dorothy B. 1979: Mushrooms of Western North America.
Univ. California Press.
Parks, H.E. 1919: Notes on California fungi. Mycologia 11: 10-21.
Peck, Charles H. 1889: Report of the New York State Museum 31: 32.
Pérez-Silva E. & E. Aguirre-Acosta 1985: Micoflora del Estado de Durango,
Mexico. Revista Mexicana de Micologia 1: 315-329.
Pilat, Albert 1936: Atlas des Champignons de l'Europe. Omphalia (Fr.) Quél.
Praga.
Redhead, S.A. & L.K. Weresub 1978: On Omphalia and Omphalina.
Mycologia 70: 556-568.
Saccardo, Pier A. 1925: Sylloge Fungorum Vol. XXIII: 46-112.
Singer, Rolf 1964: Die Gattung Gerronema. Nova Hedwigia 7: 53-92.
189
Omphalina rosella (Lange) Moser
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MY COTAXON
Volume L, pp. 191-198 January-March 1994
PHAULOMYCES SIMPLOCARIAE SP. NOV.
(ASCOMYCETES, LABOULBENIALES) FROM
SIMPLOCARIA SEMISTRIATA (COLEOPTERA,
BYRRHIDAE)
ASDEARBS BL
Nationale Plantentuin van Belgié,
Domein van Bouchout, B-1860 Meise (Belgium).
SUMMARY
A new species of the genus Phaulomyces (Ascomycetes,
Laboulbeniales), parasitic on Simplocaria semistriata Fabricius
(Coleoptera, Byrrhidae) is described and illustrated: Phaulomyces
simplocariae sp. nov.
INTRODUCTION
The genus Phaulomyces from Thaxter (1931) occurs on various
host-families and comprises, including the new species, eleven
species. Four species have been reported in Europe (Santamaria
et al, 1991; Santamaria, 1992) 1.e.: P. octotemni (T. Majewski)
I.I.Tav. on Octotemnus (Coleoptera, Ciidae) from Poland
(Majewski, 1973c); P. perparvus Santamaria on Atomaria
(Coleoptera, Cryptophagidae) from Spain (Santamaria, 1992), P.
euaestheti (Thaxt.) ILTav. on Euaesthetus (Coleoptera,
Staphylinidae) from Germany (Scheloske, 1969) and P.
denticulatus Santamaria on Phloeocharis (Scotodytes) from
Spain (Santamaria, 1992).
192
Phaulomyces simplocariae sp. nov. was found on Simplocaria
semistriata Fabricius, 1794, it is the first Laboulbeniaceous
parasite found on Byrrhidae (Coleoptera). According to
Benjamin (1989) all the previous records on Byrrhidae, i.e.
Aporomyces and Cantharomyces bordei, were of the family
Limnichidae (Limnichus, Byrrhinus, Pelochares).
The genus Phaulomyces is recognised by a two-celled primary
receptaculum bearing two or three appendages that arise from the
basal appendage cell, which is supported by cell III (Tavares,
1985). The basal appendage cell usually bears a sessile
antheridium. The perithecium has three tiers of outer wall cells
of unequal height and inconspicuous cell-walls in its basal cells.
All the specimens were mounted in Amann’s medium as
described in De Kesel (1989), they are stored in BR. The
drawings were made with a Leitz Dialux 20 EB (drawing tube).
DESCRIPTION
Phaulomyces simplocariae De Kesel sp. nov. (Figs. 1A, 1B, 1C)
Thallus hyalinus, praeter perlucidos gilvo-fuscos parietes cellularum basalium
perithecii. Fungus monoecius. Receptaculum e 3 cellulis compositum. Cellula
basalis paulo elongata, ad basim angustata. Cellula II latior quam longior,
cellulam pedunculi perithecii (cell. VI) et cellulam II’ ferens. Cellula II’ et cellula
pedunculi perithecii lateribus connatae.
Receptaculum maturum | perithecium praeditum. Cellula pedunculi perithecii
(cell. VI) isodiametra. Parietes cellularum basalium perithecii in maturis
peritheciis vix distinguibiles. Paries perithecii ex cellulis altitudine inaequalibus
compositus. Perithecium, symmetricum, fusiforme, apicem paulo differentiatum.
Cellula II’ appendicem ex cellula basalis una composita ferens, ipsa 1 (-2)
antheridiis ampuliformis et 2-4 sterilibus ramis instructa; rami 0-2 ramulis
praediti.
Longitudo a basi usque ad apicem perithecii: 95-125ym (N=61); perithecium: 60-
80pm X 16-24pm magnum (N=61); appendices ad 85-255(-400)pm _ longae
(N=61); antheridia: 11-17pm longae (N=29); ascosporae: 30pm longae (N=3).
HOLOTYPUS: ADK670 (BR), die 7.III.1993, in locis Bornemum (Antverpia,
Belgia), habitat ad sylva de Notelaria (1.F.B.L: C4.54.13), ad elytrum
Simplocariae semistriatae Fabricius (0) lectus.
i fe
ro
ez.
SOpm
nov. - 1A. Holotypus, mature thallus
Figure 1. Phaulomyces simplocariae sp
h antheridia, slide
1 wit
d thalli
Paire
with antheridium, slide ADK670. - 1B
ADK675a. - 2C. Thallus with second perithecium born on cell II’, slide
ADK666b. - Scale
mounted in Amann’s medium.
194
Thallus hyaline, except for the first tier of outer wall cells of the
perithecium, which is pale yellowish-brown. The receptacle
consists of three superposed cells. The basal cell (cell I) is
slightly elongated and becomes narrower towards the black foot.
The suprabasal cell (cell II) is broader than high and bears the
stalk cell of the perithecium (cell VI) and a somewhat larger
third receptacle cell (cell II’). Cell II’ and cell VI are connected
laterally.
The mature thallus has one, rarely two, stalked, symmetric,
fusiform perithecia with an undifferentiated apex and hardly
distinguishable basal cells. The length of the basal cells can vary
considerably (compare thalli in figure 1). The perithecial stalk
cell (cell VI) is usually isodiametric. The outer wall cells of the
perithecium are of unequal height and have inconspicuous cell
walls. The first tier is much taller than second and third together.
The usually single, relatively small isodiametric basal cell of the
primary appendage arises at the apex of cell II’. It bears two,
sometimes three or four, sterile, simple or moderately branched
appendages and one or two sessile antheridia.
Thalli with a spent or damaged perithecium can produce a
second perithecium. Old thalli usually show a precursors of a
second perithecium, i.e. procarp, on the inner or outer margins
of cell II’ (Fig. 1A).
Measurements: Total length from foot to perithecial apex: 95-
125m (N=61); Length and width of perithecium with stalk cell:
60-80pmX 16-24ym (N=61); Length antheridia: 1 1-17pm (N=29);
Length appendages 85-255ym(400pm) (n=61); Length spore:
30pm (N=3).
HOLOTYPE: Belgium (prov. Antwerpen), Bornem, Domein de
Notelaer (I.F.B.L: C4.54.13), deciduous forest, 7.[[.1993, on
Simplocaria semistriata Fabricius (0), elytrae. Leg. A. De Kesel,
(BR) ADK670.
PARATYPES: Ibid. on S. semistriata (¢) elytrae. Leg. A. De
Kesel, (BR) ADK666a, ADK666b. Ibid. on S. semistriata (&),
abdomen. Leg. A. De Kesel, (BR) ADK667. Ibid. on S.
195
semistriata (&), elytrae. Leg. A. De Kesel, (BR) ADK668a,
ADK668b. Ibid. on S. semistriata (&%), right elytron. Leg. A. De
Kesel, (BR) ADK669. Ibid. on S. semistriata (&), abdomen and
elytrae. Leg. A. De Kesel, (BR) ADK671a, ADK671b. Ibid. on
S. semistriata (&), abdomen, elytrae and femur. Leg. A. De
Kesel, (BR) ADK672a, ADK672b, ADK672c, ADK672d. Ibid.
on S. semistriata (0), abdomen. Leg. A. De Kesel, (BR)
ADK673a, ADK673b. Ibid. on S. semistriata (%), elytrae. Leg.
A. De Kesel, (BR) ADK674.
Belgium (prov. Antwerpen), Bornem, Domein de Notelaer
(sles, C4543). sdeciduous. “‘forese 2111993." one's:
semiStriata (?), abdomen and elytrae. Leg. A. De Kesel, (BR)
ADK675a, ADK675b. Ibid. on S. semistriata (%), right elytron.
Leg. A. De Kesel, (BR) ADK676. Ibid. on S. semistriata (%),
right elytron. Leg. A. De Kesel, (BR) ADK677.
DISCUSSION
The host, S. semistriata, is a widely distributed species, it occurs
in various habitats from subtropical and temperate zones (Freude
et al., 1965). In Belgium it is a common species in deciduous
forests. The adults and larvae feed on rhizoids of mosses. The
adult beetles are active from september till april (De Kesel,
unpubl. data). The beetles were captured with pitfalls. All
specimens come from a relatively humid deciduous forest near
a river. One infested specimen was found in coastal dunes.
Phaulomyces simplocariae is a relatively small species and is,
considering the hairiness of the hosts’ integuments, easily
overlooked. It usually grows in pairs on the elytrae and
abdomen. The presence of paired perithecial thalli indicates that
the fungus is monoecious.
Different infection patterns between male and female beetles
were observed, they suggest that the parasite is transmitted
during copulation. Males are mostly infested on their abdomen,
females on the caudal region of their elytrae. Statistical
196
Figure 2. Phaulomyces simplocariae sp. nov. - 2A. Immature thallus with
primary appendage and antheridium, slide ADK667. - 2B. Idem, slide
ADK672b. - 2C. Immature thallus with the perithecial initial cell formed by
cell II’, slide ADK671b. - 2D. Idem, slide ADK671b. - 2E. Immature thallus
with procarp, slide ADK675b. - 2F. Thallus with young perithecium and
trichogyne. - Scale = 50pm - all mounted in Amann’s medium.
197
treatments and more observations should be made to ascertain
this.
P. simplocariae belongs in the subtribus Euphoriomycetinae
since its mature perithecium has non persistent basal cells and
the tiers of outer wall cells are of unequal height.
The basal appendage can be considered as the upper receptacle
cell (cell III) like in Meionomyces Thaxt were cell III bears an
antheridium and the appendages. We stress that some features of
P. simplocariae indicate that Phaulomyces and Meionomyces are
strongly related. Since cell HI is not that small and a foot
extension is lacking we thought it save to place this new species
in Phaulomyces.
Some drawings were made of immature thalli (Fig. 2A, 2B, 2C,
2D, 2E, 2F), they show that the procarp is formed by the cell
above cell II, i.e. cell Il’. Wether this is the case in all other
Phaulomyces species should be studied.
Fig. 1A (Holotype) shows a lateral outgrowth on cell II’, this
outgrowth is probably a second procarp, it occurs frequently in
mature thalli. Two thalli were observed bearing a second
perithecium on the latero-apical side of cell II’ (Fig. 1C). Both
thalli clearly illustrate that a second perithecium is formed only
when the first perithecium is spent or breaks off. The possibility
that habitat characteristics or climatological factors might
influence the development of a second perithecium should be
considered.
Thalli, from the newly described species, resemble Phaulomyces
octotemni (T. Majewski) I.1.Tav. found on Octotemnus (Ciidae)
by Majewski (1973). P. simplocariae differs in two respects
from P. octotemni; the mature thallus is taller and the
perithecium is symmetrical.
ACKNOWLEDGEMENTS
Thanks are due to Dr. S. Santamaria for reviewing this paper. I
also wish to thank G. Haghebaert, for checking the determination
198
of the host. A. Fraiture is acknowledged for checking and
improving the latin description. I thank O. Van de Kerckhove for
drawing figures 1A and 1B.
REFERENCES
Benjamin, R. K. (1989). Taxonomy and morphology of
Aporomyces (Laboulbeniales). Aliso 12(2):335-367.
De Kesel, A. (1989). Ontogeny of Laboulbenia slackensis Picard
& Cépéde (Ascomycetes). Bull. Soc. Roy. Bot. Belg.
122:37-46.
Breude, “H.... ‘Harde-kW .) Lohse, GiAwe(1963)s Die {Kk ater
Mitteleuropas. Band VI. Goecke & Evers. Krefeld. 367p.
Majewski, T. (1973). Rare and new Laboulbeniales from Poland
IV. Acta Mycol. 9:229-238.
Santamaria, S. (1989). El orden Laboulbeniales (Fungi,
Ascomycotina) en la Peninsula Ibérica e Islas Baleares.
Edicions especials de la Societat Catalana de Micologia
Vol.3:1-396.
Santamaria, S. (1992). New and interesting Laboulbeniales
(Fungi, Ascomycotina) from Spain. Nova Hedwigia
54:479-492.
Santamaria, S., Balazuc, J., Tavares, I.I. (1991). Distribution of
the European Laboulbeniales (Fungi, Ascomycotina) an
annotated list of species. Treballs de l'Institut Botanic de
Barcelona vol.XIV:1-123.
Scheloske, H.W. (1969). Beitrage zur Biologie, Okologie und
Systematik der Laboulbeniales (Ascomycetes) unter
besonderer Berticksichtigung des _ Parasit-Wirt-
Verhaltnisses. Parasitol. Schriftenreihe 19:1-176.
Tavares, I.J. (1985). Laboulbeniales (Fungi, Ascomycetes).
Mycologia Memoir 9. Cramer, Braunschweig. 627p.
Thaxter, R. (1931). Contribution towards a monograph of the
Laboulbeniaceae. Part V. Mem. Amer. Acad. Arts Sci.
16:1-435.
MAY CO TAXON
Volume L, pp. 199-202 January-March 1994
STOMIOPELTIS GLOCHIDIICOLA SP. NOV.
ALAKA PANDE AND V.G.RAO
Mycology & Plant Pathology Dept.
Agharkar Research Institute, (M.A.C.S.)
Pune 411 004, India
A black, scutellate, micropeltaceous fungus was
collected on leaves of Glochidion hohenackeri Bedd. in
forests of Mahabaleshwar (M.S.) (elev. 1300 m.s.1.). The
fungus showed a non-radiate nature of the scutellum which
wasmade up of pseudoparenchymatous tissue with lobed cells,
presence of free superficial mycelium with brown, branched
hyphae, orbicular thyrothecia, bitunicate asci with
hyalodidymo- spores and septate, branched pseudoparaphyses.
All these characters showed that the collection belonged to
the genus Stomope/]tis Theiss. & Syd. emend. Luttrell.
The genus is typified by S.aspersa (Berk.)Theiss.,
collected on leaves of a species of Lauraceae from India
and described by Theissen (1914). Originally the genus was
restricted for thyrothecia with a single locule but Luttrell
(1946) emended the description to include S. polyloculata
Luttrell with several locules per thyriothecium. He
stressed that since variations in number of locules per
ascocarp occurred in a single species of the genus, the
inclusion of multiloculate species under Stomiopeltis with
Suitable expansion of generic limits was justifiable and
variation in number of locules per ascocarp should not be
considered as an important criterion for separation at
generic level. His collection on Arundinaria tetae (Walt. )
Muhl. with multiloculate thyrothecia was thus included by
him in this genus as Stomiopeltis polyloculata.
There are more than 25 species described in Stomiopel-
tis so far. Speciation is based entirely on comparative
morphology of various structures like ascocarps, asci and
ascospores.
After monographic work by Luttrell ( Luttrell, 1946),
who described the then known 7 species, several species have
been added to the genus Stomiopeltis (Muller SV LAr X
1962, Ellis, 1977).
Multiloculate thyrothecia with superficial mycelium,
non-radiate scutellum, didymosporous ascospores in the
present collection confirmed its inclusion under Stomiopel-
tis (Berk.) Theiss. emend. Luttrell.
ee
ore v. pet)
A ree
Spy]
Big eu 8 etme AF
HDG SFOS PISS
Fig.1.Stomiopeltis glochiditcola Pande & Rao
A: thyriothecium; B: a part of scutellum; C: Ascus;
D: Ascospores; E:Section of a thyriothecium.
Fig. 2.:Photomicrographs of Stomiopeltis glochidiicola Pande & Rao showing:
A: A part of the scutellum showing non-radiate, irregular cells;
B: superficial branched hyphae; C: Section of a thyriothecium
D: Asci and Ascospores.
Bar = 10 um (A,C,D); Bar = 20 um (B)
201
rable 1; Comparison of Stomiopeltis species with
multiloculate thyriothecia
Species Scutellum|Locule| Asci Ascospores
no.
Seo Case LLM
S.polyloculata |286-680 um) 2-16
Luttrell (avi V7. 25 x 4.4 um)
CL/B es. 9)
S.glochiditcola| 160-210 um| 2-5 42-50 x
ir) 8
12-15 x 3-6 um
Cavin 13. 5) C405" in)
Cia s 10)
Pande & Rao
202
After the original collection of S. aspersa by Berkeley
(Theissen, 1914) from Khasi Hills, Assam, the genus had not been
recollected from India until we collected this specimen on living
leaves of Glochidion hohenackeri Bedd. (Euphorbiaceae).
Comparison with the only multiloculate species, S.
polyloculata Luttrell, the present material showed to be
distinct. It is described here as a new species (Table 1).
Stomiopeltis glochidiicola sp. nov. Pande & Rao (Fig.1 & 2)
Mycelium superficial, composed of brown, branched
hyphae, forming minute brownish spots on leaf surface.
Ascocarps subcuticular, dimidiate, scutellate, orbicular,
160-210 um in diam., multiloculate; locules opening through
separate ostioles. Shield (scutellum) pseudoparenchymatous,
non-radiate, composed of irregularly lobed brown cells,
becoming plechtenchymatous towards margin, merging into
mycelial network. Asci pseudoparaphysate, clavate, bituni-
cate, octosporous, produced with bases towards rim and
pointing towards central ostiole, 42 - 50 x 17 - 21 um.
Ascospores hyaline, one-septate, non-constricted at septum,
rounded at both ends, 12 - 15 x 3 -— 6 um (1/b = 3.0).
Collected on living leaves of Glochidion hohenackerit
Bedd. at Mahabaleshwar (M.S., India) (el. 1300. Msel
dt. 6.1.1982, AMH 7149 (Holotype).
Maculae brunneae, orbiculares. Mycelium superficiale.
Ascomata subcuticularia, 160 - 210 um diam., multilocu-
lata. Loculae 2 - 5, ostiolatae. Scutellum pseudoparenchyma-
ticum, non-radiatum, ex celluis brunneis' irregulariter
lobatis, marginam versus ex cellulis plectenchymatibus
formatum. Asci pseudoparaphysatibus, clavati, bitunicati,
octospori, 42 -50 x 17 - 21 um. Ascosporae hyalinae, unisep-
tatae, non-constrictae, ad apicem rotundatae, 12 - 15 x 3 -
6 wm.
In foliis viventibus Glochidit hohenackeri Bedd.,
loco: Mahabaleshwar (M.S., India), dt. 6.Jan.1982, holotypus
lectus ad AMH subnumero 7149.
Authors thank Dr. B.C, Sutton, I.M.I., England, for
suggestions on the manuscript.
References
Ellis, J. P. 1977. The genus Stomiopeltis in Britain.
Trans. Br. mycol. Soc. 68(2): 157-159.
Luttrell, E.S. 1946. The genus Stomiopeltis (Hemisphaeria-
ceae) Mycologia 38: 565-586
MUller, E. & J.A.von Arx 1962. Die Gattungen der didymospo-
ren Ascomyceten. Beih. zur Kryptogamenfl. der
SCOW = elena oe...
MY COTAXON
Je tea Th A A EE he dE he BL aR ARE
Volume L, pp. 203-217 January-March 1994
ADDITIONAL NEW SPECIES AND NEW REPORTS OF PERTUSARIA
(LICHENISED ASCOMY COTINA) FROM NEW ZEALAND WITH A REVISED
KEY TO THE CORTICOLOUS SPECIES IN NEW ZEALAND.
ALAN W. ARCHER
National Herbarium of New South Wales,
Royal Botanic Gardens, Sydney, NSW, Australia, 2000
JOHN A. ELIX
Department of Chemistry, The Faculties, Australian National University,
Canberra, ACT, 0200, Australia
ABSTRACT: The species Pertusaria barlettii Archer & Elix, Pertusaria celata
Archer & Elix, Pertusaria hadrospora Archer & Elix, Pertusaria parvula Archer &
Elix, Pertusaria psoromica Archer & Elix, Pertusaria scutellifera Archer & Elix
and Pertusaria sporellula Archer & Elix are described as new; Pertusuria
leucoplaca Mill.Arg., Pertusaria schizostomella Mill.Arg., Pertusaria
subisidiosa Archer, Pertusaria subplanaica Archer & Elix, Pertusaria thamnolica
Archer and Pertusaria velata (Turn.) Nyl. are reported as new to New Zealand.
Pertusaria albissima Miill.Arg. and Pertusuria alboatra Zahlbr. are re-instated into
the New Zealand lichen flora. A key to the New Zealand corticolous species of
Pertusaria is given.
INTRODUCTION
The lichen genus Pertusuria in New Zealand was reviewed by Galloway (1985) and an
account of the saxicolous species common to Australia and New Zealand has been given
(Archer & Elix 1993). An examination of additional recent corticolous specimens from
New Zealand |from AK, AKU, CHR, OTA and WELT] has shown the presence of both
undescribed taxa and species new to New Zealand and these are reported here. The
techniques used for the examination of specimens have been described previously
(Archer 1991); the chemistry of the majority of the specimens examined was confirmed
by liquid chromatography (Elix & Venables 1993). The spore descriptions follow the
nomenclature of Dibben (Dibben 1980: 10). The figures illustrate holotypes and, unless
noted otherwise, the type specimens cited have been seen. A key to the New Zealand
corticolous species of Pertusaria is given.
Pertusaria bartlettii Archer & Elix sp. nov. Fig. |
Thallus flavido-albus, areolatus et rimosus, superficies laevis et hebetata, isidiis et
sorediis destitutis; corticola; apothecia verruciformia, inconspicua, numerosa, flavida,
saepe confluentia, perplano-hemisphaerica vel immersescia, 1-2 mm diam., ostiola
conspicua, nigra, circularia vel irregularia, 0.2-0.6 mm latae, in verrucas |-3na; sporae
8nae, biseriatae, ellipsoideae, laeves, 60-75 wm longae, 25-32 jam latae.
Thallus arthothelin, acidum thiophanicum et 2,4-dichloronorlichexanthone continens.
204
Type: New Zealand, North Island, Mangonui County, Herekino Gorge, 35°10'S,
173°12' E, alt. 400 ft. [ca.120 m], on bark of Rhopalostylus sapida on exposed hillside,
J.K. Bartlett s.n., 26.x.1981; AK 192225-holotype.
Thallus pale yellowish white, areolate and cracked, surface smooth and dull, lacking
isidia and soredia, corticolous; apothecia verruciform, inconspicuous, numerous, pale
yellow, sometimes confluent, very flattened-hemispherical or becoming immersed, 1-2
mm diam.; ostioles conspicuous, black, circular or irregular in outline, 0.2-0.6 mm wide,
]-3 per verruca; spores 8 per ascus, biseriate, ellipsoid to subfusiform, smooth, 60-75
pm long, 25-32 wm wide.
Chemistry: K-, KC+ orange red, C+ orange, Pd-; arthothelin (major), thiophanic acid
(major), 2,4-dichloronorlichexanthone (major ); 2,5-dichloronorlichexanthone (minor)
and 4,5-dichloronorlichexanthone (trace), 2-chloronorlichexanthone (trace), 4
chlorolichexanthone (trace) and thiophaninic acid (trace).
Pertusaria bartlettii is characterised by pale yellow, flattened or immersed verrucae with
conspicuous black ostioles and the presence of polychlorinated norlichexanthones; these
features distinguish the new species from all other New Zealand Pertusaria.
The new species is known only from the type specimen; the epithet "bartlettii" refers to
John K. Bartlett, 1945 -1986, an enthusiastic collector of lichens in New Zealand.
Pertusaria celata Archer & Elix sp. nov. Figs?
Thallus albidus vel cinereo-albidus, tenuis, rimosus, superficies laevis et nitidae, isidtis et
sorediis destitutis; corticola; apothecia verruciformia, conspicua, sparsa, plano-
hemisphaerica, thallo concoloria, basibus non constrictis, lacerescentia, epithecia fusca
vel nigra manifestescentia; sporae 8nae, biseriatae, ellipsoideae vel subfusiformes,
laeves, 90-100(-120) wm longae, 30-40(-50) zm latae, parietibus exterioribus 2-4 wm
crassis, parietibus interioribus 4-5 jm crassis.
Thallus acida salazinicum, norsticticum et connorsticticum continens.
Type: New Zealand, North Island, South Auckland, Matamata County, Mamaku Plateau,
Tirau-Matamata Road, 38°09'S, 176°E, alt. 1200 ft |ca. 360 m], on Weinmannia
racemosa bark, J.K. Bartlett 19749, 28.11.1982; AK 192245-holotype.
Thallus off-white to pale grey, thin, cracked, surface smooth and shiny, lacking isidia
and soredia, corticolous; apothecia verruciform, conspicuous, sparse, flattened-
hemispherical, concolorous with the thallus, not constricted at the base, 1-1.5 mm diam.,
becoming split and torn to reveal the dark brown to black epithecium; spores 8 per ascus,
biseriate, ellipsoid to subfusiform, smooth, 90-100(-120) pm long, 30-40(-50) wm wide,
outer wall 2-4 wm thick, inner wall 4-5 ym thick.
Chemistry: K+ yellow becoming red, KC-, C-, Pd+ yellow; salazinic acid (major),
norstictic acid (minor) and connorstictic acid (trace).
Specimens examined:
New Zealand, North Island, type locality, , J.K. Bartlett 19739, 28.11.1982 (AK
192244), J.K. Bartlett 19736, 28.111.1982(AK 192246).
South Island, Nelson, Mt. Duppa Track, 41°12'S, 173°29'E, alt. ca. 750 m, W. Malcolm
855, 3.v.1993 (CHR 470255).
Pertusaria celata is characterised by the torn verrucae, revealing the epithecium of the
apothecia below and in this respect it somewhat resembles the saxicolous New Zealand
205
Figs 1-7. New Pertusaria species. 1. Pertusaria bartlettii, x3; 2. Pertusaria celata, x3;
3. Pertusaria hadrospora, x4; 4. Pertusaria parvula, X3.75; 5. Pertusaria
psoromica, x3; 6. Pertusaria scutellifera, x3; 7. Pertusaria sporellula, x4.
206
P. erumpescens Nyl.; the spores in this species are 30-40 ym long. Although unlike
many verruciform Pertusaria taxa, the new taxon has spores with the double wall
characteristic of the genus. P. celata is known from both the North and South Islands.
The epithet "celata" hidden, refers to the concealed epithecium and asci.
Pertusaria hadrospora Archer & Elix sp. nov. Fig. 3
Thallus dilute olivaceus, crassus, subrimosus, superficies laevis et nitidae, isidiis et
sorediis destitutis; corticola; apothecia verruciformia, inconspicua, saepe solum ostiolis
indicata, perplano-hemisphaerica vel immersa, thallo concoloria, 0.6-1 mm diam.; ostiola
inconspicua, subtranslucida, in verrucas |-2na, 0.2-0.4 mm diam.; sporae 2nae, raro 3,
ellipsoideae, laeves, 210-250(-300) wm longae, 50-75 jm latae.
Materia chemica in thallo non detecta.
Type: New Zealand, South Island, Golden Bay County, northwest Nelson, Kaihoka
Lakes, 40°31'S, 172°36'E, sea-level, J.K. Bartlett 22268, 13.xii.1982; AK 192257-
holotype.
Thallus pale olive green, thick, somewhat cracked, surface smooth and shiny, lacking
isidia and soredia, corticolous; apothecia verruciform, inconspicuous, often only
indicated by the ostiole, very flattened-hemispherical or immersed, concolorous with the
thallus, 0.6-1 mm diam.; ostioles inconspicuous, grey, translucent, lor 2 per verruca,
0.2-0.4 mm diam.; spores 2 per ascus, rarely 3, ellipsoid, smooth, 210-250(-300) pm
long, 50-75 wm wide.
Chemistry: K-, KC-, C-, Pd-; no lichen compounds detected by tle or hple.
Specimen examined:
New Zealand, South Island, Golden Bay County, northwest Nelson, Tasman
Mountains, Lake Cobb, 41°02'S, 172°31'E, alt. ca 3000 ft.[900 m], J.K. Bartlett 2/528,
14.xii. 1982 (AK 192258); Arthur Range, Lodestone track, 41°10.7'S, 172°44.6'E, alt.
1170 m, W. Malcolm 992, 18.vii.1993 (herb. Malcolm).
Pertusaria hadrospora is characterised by the inconspicuous verrucae and asci with two
(rarely three) large spores; these features distinguish the new species from all other
corticolous New Zealand Pertusaria.
Zahlbruckner (1941: 334) reported a corticolous, two-spored Pertusaria, P.tetrathalumia
var. major |P. leucoplacoides var. major Miill.Arg. (Miller 1884: 301)]| to occur in
New Zealand but the specimen on which this report was based (CHR 452614) is
identified as P. sorodes Stirton. The spores in the specimen are 225-260 ym long and 55-
70 xm wide, with rough inner spore walls, whereas Miiller reported the spores in his
var. major to be 150-210 wm long and 30-40 ym wide (Miiller, loc. cit.). This specimen
is morphologically and chemically distinct from P. hadrospora.
The epithet "hadrospora", well-devloped spores, refers to the large spores present in the
asci of the new species.
Pertusaria parvula Archer & Elix sp. nov. Fig. 4
Thallus albidus, tenuis, subrimosus, superficies subnitida, isidiis et sorediis destitutus;
corticola; apothecia verruciformia, perplano-hemisphaerica, thallo concoloria, dispersa,
0.8-1.2 mm diam.; ostiola conspicua, nigra, 0.2-0.3 mm diam., in verrucas singula;
sporae 8nae, uniseriatae, ellipsoideae, laeves, 30-37 wm longae, 18-20 ym latae.
Thallus 4,5-dichlorolichexanthone solum continens.
207
Type: New Zealand, South Island, Sounds-Wellington Ecological Region, Cook Strait
Ecological District, Chetwode Islands, Nukuwaitata Island, 40°54'S, 174°04'E, on bark,
B. Hayward, s.n., Mar. 1984; AK 206954-holotype.
Thallus off-white, thin, somewhat cracked, surface smooth and somewhat shiny, lacking
isidia and soredia; corticolous; apothecia verruciform, very flattened hemispherical,
concolorous with the thallus, scattered, 0.8-1.2 mm diam.; ostioles conspicuous, black,
0.2-0.3 mm diam., | per verruca; spores 8 per ascus, uniseriate, ellipsoid, smooth, 30-
37 pm long, 18-20 pm wide.
Chemistry: K-, KC-, C-, Pd-; 4,5-dichlorolichexanthone only.
Pertusaria parvula is characterised by verruciform apothecia, small spores and the
presence of 4,5-dichlorolichexanthone. Externally it resembles both P. melaleucoides
Miill.Arg. and P. albissima Mill.Arg. but is distinguished from those species by the
number and size of spores, and the presence of the chlorinated xanthone. The new
species is known only from the type specimen.
The epithet "parvula" (smallest) refers to the small spores characteristic of the new
species.
Pertusaria psoromica Archer & Elix sp. nov. Fig. 5
Thallus hinnuleus vel dilute olivaceus, areolatus et rimosus, superficies laevis vel
subtuberculata, subnitida, isidiis destitutis sed sorediatus; corticola; soralia conspicua,
numerosa, alba, hemisphaerica vel disciformia, 0.5-2 mm diam.; apothecia non visa.
Thallus acidum psoromicum continens.
Type: New Zealand, North Island, Otamatea County, Kaiwaka, 36°09'S, 174°27'E, J.K.
Bartlett 24219, 1975-1985, AK 19223 |-holotype.
Thallus dull fawn or pale olive green, areolate and cracked, surface smooth or
subtubercular, somewhat shiny, lacking isidia, sorediate, corticolous; soralia
conspicuous, numerous, white, initially pustulate, composed of granular soredia,
hemispherical or discoid, 0.5-2 mm diam.; apothecia not seen.
Chemistry: K-, KC+ yellow, C+ yellow, Pd+ strong yellow; psoromic acid (major) and
conpsoromic acid (minor).
Specimens examined:
AUSTRALIA, Queensland, Mt. Baldy, 4 km SW of Atherton, 17°17'S, 145°27'E, alt.
1080 m, on Alphitonia in rain forest, J.A.Elix 16276, 25.vi.1984 (ANUC); Mount
Windsor Tableland, 45 km NW of Mossman, 16°15'S, 145°OI'E, alt. 1200 m, on
Syzygium in rain forest, J.A.Elix 16445, 16449, 26.vi.1984 (ANUC).
New South Wales, Wiangaree Forest Drive, Tweed Range, "Antarctic Beech Walk", on
Nothofagus moorei in rain forest, G. Kantvilas 645/88, 3.viii.1988 (HO, NSW
219573); Border Ranges National Park, Bar Mountain Lookout track, 28°28'S,
153°06'E, alt. ca. 950 m, on fallen branch, A.W. Archer P 480, 7.1x.1992 (NSW).
NEW ZEALAND [15 out of 39], North Island, Great Barrier Island, Mount Hobson,
36°11'S, 172°2S'E, alt. ca. 500 m, B.Hayward s.n., Jan. 1983 (AK 182649); ibid., Jan.
1984 (AK 171774); Rakitu Island, 36°08'S, 175°30'E, B.Hayward s.n., 1980
(AK162392); ibid. (AK 160765); Coromandel County, Mount Moehau, 36°32'S,
175°24'E,alt. ca. 700 m, B. Hayward H39.65, Aug. 1974 (AK 154517); Thames
County, Upper Kauaeranga Gorge, 37°04'S, 175°40'E, alt. ca. 350 m, J.K.Bartlett s.n.,
1975-1985 (AK 192243); Rimutaka State Forest Park, 41°21'S, 174°5S8'E, alt. 300 m, on
Pseudowintera colorata in Kunzea ericioides forest, B. Polly s.n., 1.vi.1993 (WELT).
208
South Island, Dunedin, Mt. Cargill, ca. 300 m, on bark of Aristotelia, W.Martin s.n.,
9.xi1.1953 (OTA 880); Westland, between Greymouth and Runanga, on Nothofugus, W.
Martin s.n., 4.11.1954 (OTA 1285); Otago, Silver Peaks near Dunedin, on beech, J.
Thomson 1229 (CHR361434); West Otago, Eglington Valley, Cascade Creek, on
Nothofagus menziesii, alt. ca. 450 m, W.Martin s.n., 27.11.1972 (CHR479324 pp);
Golden Bay, Kaihoka, 40°33'S, 172°3S'E, J.K. Bartlett s.n. (AK 192253, AK 192254);
Stewart Island, Port Pegasus, Islet Cove, 47°12'S, 167°38’E, alt. 10 m, B.- Hayward
s.n., 3.11.1989 (AK208187); D'Urville Island, Mount Maud, 40°53'S, 173°49'E, alt. ca.
500 m, B.Hayward s.n., 4.1.1988 (AK206939).
Pertusaria psoromica 1s characterised by the presence of numerous, conspicuous white
soralia, reacting K- and Pd+strong yellow due to the presence of psoromic acid. The
soredia of P. psoromica also give a yellow colour with KC and C; this colour is
produced by reaction with psoromic or conpsoromic acids as thin layer chromatography
showed only these two compounds to be present. The podetia of Cladonia subpityrea
Sandst. (which also contain the same two lichen acids) give no colour with KC and C but
if acetone extracts from both species are spotted onto filter paper and treated with KC or
C, a yellow colour is given by both extracts. The new species occurs at all altitudes from
sea-level to cu. 700 m in New Zealand and up to 1200 m in Australia. In New Zealand it
has been found on Coprosma arborea, Nothofagus menziesii, Olearia furfuracea, O. rani,
and on the bark of Rhopalostylis sapida, Metrosideros excelsa and Dysoxylum
spectabile. The new species resembles, in morphology and wide distribution, the sterile,
sorediate Australian P. scaberula Archer (Archer 1991:240) but is distinguished from that
species (which contains lichexanthone and thamnolic acid) by the negative reaction with
alkali. Pertusuria scaberula is common in eastern Australia whereas P. psoromica occurs
in both the North and South Islands of New Zealand, and to a more limited extent in
Australia, in Queensland and New South Wales.
The new species is chemically similar to P. sphaerulifera Vainio (Vainio 1907)
from Thailand but that species is saxicolous, and pustulate rather than sorediate.
Pertusaria scutellifera Archer & Elix sp. nov. Fig. 6
Thallus albidus, areolatus et mmosus, superficies irregulariter subtuberculata et hebetata,
isidiis et sorediis destitutis; corticola; apothecia disciformia, conspicua, numerosa, thallo
concoloria, 0.6-1.2 mm diam.; disci auranto-fusci et confertim albo-pruinosi; sporae
singulae, ellipsoideae, laeves, (125-)150-185 um longae, 45-55 jum latae, parietibus 3-5
pam Crassis
Thallus acidicum thamnolicum continens.
Type: New Zealand, North Island, Mt. Ruapehu, W.H.Ewers 2657, 31.ii1.1988; AK
200760-holotype.
Thallus off-white, areolate and cracked, surface irregularly subtuberculate and dull,
lacking isidia and soredia; corticlous; apothecia disciform, conspicuous, numerous and
crowded, concolorous with the thallus, 0.6-1.2 mm diam.; disc orange-brown, densely
white-pruinose; spores | per ascus, ellipsoid, smooth, (125-)150-185 jm long, 45-55
pm wide; spore wall single, 3-5 wm thick.
Chemistry: K+ yellow, KC-, C-, Pd+ yellow; thamnolic acid.
Specimens examined:
New Zealand, North Island, Mt. Ruapehu, W.H.Ewers 2626 p.p., 2627, 31.11.1988
(CBG); Rangitaiki, W.H.Ewers 286/, 28.111.1988 (CBG).
South Island, Arthurs Pass, on Nothofagus cliffortioides, H.H.Allan s.n. (CHR
361433); West Otago, Eglington Valley, Cascade Creek, W. Martin A552, 14.11.1964
(CHR479323): Nelson, Mt. Arthur, 41°12'S, 172°44'E, 1170 m, on Dracophyllum
traversii bark in montane low forest, P.N. Johnson 4/7, 29.1.1993 (CHR 480406);
209
Nelson, Mt. Arthur, Flora Hut, start of track, 41°11'S, 172°43’'E, alt. 940 m, on fallen
branch, W. Malcolm 873, 13.v.1993 (CHR 470257).
Pertusaria scutellifera is characterised by disciform apothecia, single spored asci and the
presence of thamnolic acid. It is morphologically similar to P.novaezelandiae Szatala but
is distinguished from that species by the yellow colour given with alkali, in contrast to the
violet colour given by P. novuezelandiae which contains hypothamnolic acid. The new
species also resembles the Japanese P. sphaerophora Oshio (Oshio 1968) but is
distinguised from that species by differences in chemistry and spore morphology. Oshio
(loc.cit.) reported that with alkali (K) his new species gave a yellow colour becoming
violet and Dibben (1980) found thamnolic acid and an unidentified compound present in
the Japanese species. The two species also differ in spore wall thickness; the spore walls
are 13-21 yum thick in P. sphaerophora and 3-5 ym thick in P. scutellifera. The new
species is differentiated from the Japanese P. nigrodisca Oshio (Oshio 1968) [which also
contains thamnolic acid and one-spored asci] by the colour of the discs, black in P.
nirodisca and orange brown in P.scutellifera.
Zahlbruckner identified (in sched.) the specimen CHR 361433 as P. globulifera subsp.
glaucomopsis Nyl. [P. globulifera *glaucomopsis Nyl. tax. vag. (Nylander 1888: 67)|
but the relevant specimens in the Nylander Herbarium [H-NYL 23765, 23766, 23767,
23768] lacked thamnolic acid and this taxon is thus distinct from P. scutellifera.
Pertusaria globulifera *glaucomopsis was reported by Nylander (loc.cit.) to be K -ve.
The epithet "scutellifera", bearing little dishes, refers to the numerous conspicuous
disciform apothecia.
Pertusaria sporellula Archer & Elix sp. nov. Fig. 7
Thallus albidus vel cinereo-albidus, tenuis, superficies laevis et subnitida, isidiis et
sorediis destitutis; corticola; apothecia disciformia, numerosa,dispersa, saepe contigua,
marginibus incrassatis, initio involutis, thallo concoloribus; disci nigri, albo-pruinosi,
concavi complanatescentes, 0.2-0.4 mm diam.; sporae 8nae, uniseriatae, sphaericae, |5-
20 ym diam., parietibus ca. | yum crassis.
Materia chemica in thallo non detecta.
Type: New Zealand, North Island, Kaipera Ecological Region & District, North Kaipera
Barrier, Tapu Bush, 36°19'S, 174°04'E, alt. 110 m, A.E.Wright //56/, 25.v.1991; AK
202378-holotype.
Thallus off-white to pale greyish white,thin, surface smooth and somewhat shiny,
lacking isidia and soredia; corticolous; apotheciu disciform, numerous, scattered or
sometimes contiguous, margins thick, initially inrolled, concolorous with the thallus,
discs black, partly white-pruinose, initially concave, becoming flat, 0.2-0.4 mm diam.;
spores 8 per ascus, uniseriate, spherical, 15-20 zm diam., spore wall thin, ca. | wm
thick.
Chemistry: K-, KC-, C-, Pd-; no lichen compounds detected.
Pertusaria sporellula is characterised by disciform apothecia, eight-spored asci with very
small spherical spores and the absence of lichen compounds. It somewhat resembles P.
truncata Krempelh. but is readily differentiated from that taxon by the negative reaction
with alkaline hypochlorite (KC), in contrast to the characteristic violet colour given by
P.truncata.
The epithet "sporellula", very small spores, refers to the small spherical spores in the new
species.
210
NEW REPORTS
Pertusaria albissima Miill.Arg. Flora 67: 350 (1884)
Type: New Zealand, sin. loc., C. Knight s.n., "Pertusaria leucodeoides Knight" nom.
nud.; G-lectotype [here selected]; C. Knight s.n., "Pertusaria leucodes var. inconspicua
Knight" nom. nud.; G residual syntype.
Thallus chalky white, slightly areolate and cracked, surface smooth and dull, lacking
isidia and soredia, corticolous; apothecia verruciform, concolorous with the thallus,
conspicuous, numerous, sometimes confluent, flattened-hemispherical, sometimes
constricted at the base, 1-2 mm diam.; ostioles inconspicuous, pale, translucent, 2-4 per
verruca; spores 8 per ascus, uniseriate, ellipsoid, smooth, 50-62 wm long, 23-28 pm
wide.
Chemistry: K-, KC-, C-, Pd-; no lichen compounds detected by hplc.
Miiller described three species from different parts of Knight's specimen of "Pertusaria
leucodeoides Knight" [nom. nud. in sched.| (two of which are here reported to be
synonyms) as shown in Table 1.
Galloway (1985) reported P. albissima to be synonymous with P. cretacea
Miill.Arg. but the two species are distinguished by the difference in spore arrangement
and the colour of the ostioles (black in P. cretacea , syn. P. leucodes Knight) as
originally noted by Miiller (1884, loc.cit.) and the two species are here retained as
separate entities. Pertusaria laevis Knight is distinguished from P. albissima by the black
ostioles and the presence of thiophaninic acid, which was detected in the holotype
(WELT) by liquid chromatography. Pertusaria albissima is known only from Knight's
two specimens.
The protologue to P. albissima refers to two syntypes collected by Charles
Knight; one of these, labelled "P. leucodeoides Knight" by Knight, is also labelled "P.
albissima Miill.Arg. L.B.n. 749", a reference to the publication Lichenologische Beitrage
XIX, number 749 (Miiller, 1884) and this specimen is here selected as lectotype.
TABLE 1
NEW SPECIES FROM "PERTUSARIA LEUCODEOIDES KNIGHT"
Original Published spore spore Chemistry
Herbarium name Name arrangement length
pm
P. leucodeoides Knight — P. leucodeoides Miill.Arg. 2-seriate 90-113 xanthone(a)
|syn. P. theochroa Kremplh. 2-seriate 87-113 thiophaninic
acid (b)]
P. cretacea Miill.Arg. 2-seriate 50-70 nil (a)
[syn. P. leucodes Knight 2-seriate 55-70 nil (b)]
P. albissima Mill.Arg. l-seriate 50-62 nil (b)
P. leucodes var.
inconspicua Knight * l-seriate 48-55 nil (b)
(a) reported by Galloway, op. cit. (b) determined by Archer & Elix
ZirT
Both P. leucodeoides Miill.Arg. and P. theochroa Krempelh. (Krempelhuber
1876: 452) have verrucae with black ostioles and asci with 8 biseriate spores that are 90-
113 ym long. Galloway (Galloway 1985: 375,380) reported the chemistry of the two
taxa to be identical; the holotype of P. theochroa (M) contains thiophaninic acid. As the
two species are morphologically and chemically identical, the two species are considered
to be conspecific, as was originally suggested by Galloway (Galloway, loc. cit.).
In addition, both P. cretacea Miill.Arg. and P. leucodes Knight have abundantly
fertile thalli with numerous verrucae, the verrucae with inconspicuous ostioles and the
asci with 8 biseriate spores that are 50-70 ym long. Pertusaria leucodes was reported to
contain stictic acid (Galloway 1985: 376) but an examination of the lectotype from WELT
found no lichen compounds present. In view of these morphological and chemical
similarities, the two species are considered to be identical, as was originally proposed by
Dibben (Dibben 1984, in sched.).
Pertusaria alboatra Zahlbr. Denkschr. Akad. Wiss. Wien math.-naturwiss. Kl.
104:336 (1941).
Type: New Zealand, North Island, Rangitoto Island, Auckland, on trunk of Avicennia
officianalis in salt lagoon, H.H.Allen 46, Aug.1933; W-lectotype [fide D.J. Galloway
26.xi1.1981, in sched.]; CHR 374706-iso-lectotype; ibid., on Avicennia resinifera, H.H.
Allen W93, 12.1v.1936; CHR 361450, W-residual syntypes.
Thallus pale yellowish fawn, conspicuously areolate and cracked, surface smooth and
dull, lacking isidia and soredia, corticolous; apothecia verruciform, numerous,
conspicuous, often confluent, flattened-hemispherical to immersed, 0.5-1 mm diam.;
ostioles conspicuous, black, one per verruca, 0.1-0.4 mm diam., sometimes with a pale
grey translucent margin; spores 2 per ascus, rarely 3, ellipsoid, smooth, (70-)85- 1 10(-
132) pm long, 35-45 ym wide.
Chemistry: K-, KC+ weak yellow, C-, Pd-; thiophaninic acid (major) with 2-chloro-6-O-
methylnorlichexanthone (trace) and 4-chloro-6-O-methylnorlichexanthone (trace).
Specimens examined:
NEW ZEALAND, North Island, Mangonui County, Waitiki Stream, 34°30'S, 172°S0'E,
J.K. Bartlett s.n., 1975-1985 (AK 194384); North Island, Paihia, on mangroves, W. H.
Ewers 2743, 6.iv.1988 (CBG); Sounds- Wellington Ecological Region, Sounds
Ecological District, Resolution Bay, E end of Bay E of wharf, 41°07'S, 174°13'E, alt. 1-
5m, on bark of karaka and mahoe, B.W. Hayward s.n., 6.1.1992 (AK 205603).
Pertusaria alboatra was not included in the Flora of New Zealand Lichens (Galloway
1985) and a description has therefore been given here. The species resembles P.
melaleucoides Miill.Arg. but is distinguished from that species by the presence of
thiophaninic acid and the occasional occurrence of three-spored asci. Zahlbruckner
labelled two specimens, CHR 374706 and AK 18997, "Pertusaria atroalba (sic) sp.
nov." The specimen in CHR is numbered "46" and was collected by W.H.Allen on
24. viii. 1933 from Avicennia officinalis and contains thiophaninic acid; it is thus identical
to the specimen in W. The specimen in AK was collected at the same location on the same
day but from A. resinifera; it has no number and contains no lichen compounds and is
identified as P. melaleucoides Miill.Arg. (Archer & Elix 1993a).
Pertusaria leucoplaca Miill.Arg. Flora 67: 304 (1884).
Type: Brazil, Apiahy, on Araucaria, Puiggari 2128, Oct. 1882; holo: G.
Pertusaria acuta Miill.Arg. Flora 67: 353 (1884).
212
Type: India, in montibus Neilgherries [Nilgiri Hills, ca. 250 km SSW of Bangalore], on
Cinchona succirubris, Bader s.n.; holo: G.
Thallus greyish-white, areolate and cracked, surface smooth and dull, lacking isidia and
soredia, corticolous; apothecia verruciform, conspicuous, scattered, flattened
hemispherical, sometimes constricted at the base, concolorous with the thallus,
sometimes slightly concave above, 0.8-1.5 mm diam.; ostioles conspicuous, pale brown
to dark brown to black, 1, rarely 2, per verruca; spores 8 per ascus, biseriate or
irregularly biseriate, subfusiform to fusiform, smooth, 70-90, rarely tol100 ym long, 25-
35 ym wide.
Chemistry: K-, KC-, C-, Pd-; 2-O-methylstenosporic acid with 2-O-methylperlatolic and
2-O-methyldivaricatic acid as minor or trace compounds.
Specimens examined:
AUSTRALIA, New South Wales, Upper Kangeroo River Rd, 9 km S of Robertson, H.
Streimann 35796 (B, CBG); Olney State Forest, ca. 25 km W of Morisset, A. Archer P
166, 21.vi.1991 (NSW); Mt. Ebsworth Rd, 4 km SW of Stroud, A. Archer P194,
31.111. 1992 (NSW ); Murramarang N.P. ca. 10 km N of Batemans Bay, A. Archer P30]
(NSW ). Victoria, Laughton Gully, Mar 1889, F. Wilson (NSW).
NEW ZEALAND, North Island, Great Barrier Island, Rakitu (Arid) Island, 36°08'S,
175°30'E, B.Hayward s.n., Jan.1981 (AK169175); Poor Knights Island, Aorangi, E of
Oneho Hill, 35°30'S, 174°33' E, alt. 180 m, B.Hayward s.n., ‘Aug. 1984 (AK172444).
P. leucoplaca resembles the Australian species P. leucostigma Mill.Arg. (with which it
often occurs in Australia) but is distinguished from that species by the absence of
lichexanthone,the darker ostioles and the slightly smaller biseriate spores. Pertusaria
leucoplaca is characterised by the eight biseriate spores and the presence of 2-O-
methylstenosporic acid. It occurs in Australia, New Zealand, Brazil and India.
Pertusaria schizostomella Miill.Arg. Bull. Herb. Boissier 3: 637 (1895).
Type: Australia, New South Wales, C. Knight 3/1, 1887; G-holotype.
Thallus pale yellowish green to pale greyish green, slightly or coarsely areolate and
cracked, surface smooth or very slightly rough, dull, lacking isidia and soredia,
corticolous; apothecia verruciform, conspicuous, scattered or crowded and becoming
confluent, concolorous with the thallus, slightly flattened-hemispherical, not constricted
at the base, 0.5-1 mm diam.; ostioles conspicuous, pale, translucent, dull yellow, usually
one per verruca; spores 2 per ascus, ellipsoid, smooth, or very slightly rough, 80-125
pm long, 30-45 pm wide.
Chemistry: K-, KC+ orange, C+ orange, Pd-; thiophaninic acid (major), stictic acid
(major) with constictic acid (minor -trace).
Specimens examined:
AUSTRALIA, Queensland, Turkey, on Rhizophora, N. Stevens 1241, 18.viti.1975
(BRI); Brisbane area, Deception Bay, on mangrove, H.7. Lumbsch 5381, 10.viii.1987
(herb. Lumbsch). New South Wales, Lord Howe Island, Valley of Shadows,
31°31'45"S, 159°04'45"E, alt. 40m, J.A. Elix 32840, 3284], 22.vi.1992 (ANUC);
Newport, on mangroves, F.R.M. Wilson s.n., Oct 1888, (NSW L4499); Batemans Bay,
35°44'S, 150°13'E, sea-level, on Avicennia, H. Streimann 37917, 20.1.1987 (CBG
9201737).
213
NORFOLK ISLAND, Ball Bay Reserve, 29°32'S, 167°59'E, alt. 80 m, on Araucaria
trunk, H. Streimann 31726, |.xii.1984 (CBG 9201715); Philip Island, Upper Long
Valley, 29°07'30"S, 167°57'E, alt. 80 m, on roots of Lagunaria, H. Streimann 32202,
4.xii. 1984 (CBG 9215716); ibid., H. Streimann 32217, 4.xii.1984, (CBG 9201717).
NEW ZEALAND, North Island, Rangitoto Island, near Islington Bay, on Avicennia
resinifera, H.H. Allan s.n., 24.viii.1933 (AK 18993); ibid., 12.iv.1936 (CHR 413815);
ibid., on Avicennia officinalis, H.H.Allan s.n., 24.viii.1932 (CHR 452611, CHR
452613); near Slipper island, Watchman Rock, ST OSS; Wane ESB: Hayward 126,
Aug. 1973 (AK 171693); Coromandel County, Great Mercury Island, 36°36'S,
175°48'E, B. Hayward H40.48, May 1975 (AK 187603); Tairua Ecological District,
Slipper Island, 37°03'S, 175°56'E, B. Hayward H49.170 , H49.204, Aug. 1973 (AK
206948, AK 155028); Tapotupotu Bay, 34°36'S, 172°43'E, on mangroves, B. Hayward
s.n., May 1984 (AK 206952).
Pertusaria schizostomella is characterised by its two-spored asci, conspicuous translucent
ostioles and the presence of thiophaninic and stictic acids. It occurs in eastern coastal
Queensland and New South Wales, Norfolk Island, Lord Howe Island and in the North
Island of New Zealand, often on mangroves at sea-level.
The earliest New Zealand specimens, AK 18993, CHR 413815, CHR 45261 1
and CHR 452613, were originally identified by Zahlbruckner as P. melaleuca var.
heterochroa (Mill.Arg.) Stirton (Zahlbruckner 1941: 329) but the specimens correspond
in all respects to P. schizostomella.
Pertusaria subisidiosa Archer, Mycotaxon 41: 242 (1991).
Type: Australia, Queensland, North Stradbroke Island, NE of Brown Lake, 27°29'S,
153°26'E, on fallen trunk in mixed rain forest, J. Hafeliner 19204, 10.viii.1986; GZU-
holotype.
Thallus pale cream-white,thin, continuous, surface smooth and dull, isidiate, lacking
soredia,; corticolous; isidia simple, concolorous with the thallus, profuse, 0.1-0.3 mm
tall, 0.05-0.1 mm diam.; apothecia verruciform, concolorous with the thallus,
inconspicuous, scattered, flattened-hemispherical, isidiate, 0.5-0.7 mm diam.,; ostioles
black, conspicuous, 0.1-0.2 mm diam., | per verruca; spores 4 per ascus, uniseriate,
ellipsoid, rough, 80-95 ym long, 25-35 wm wide.
Chemistry: K-, KC-, C-, Pd-; 2,5-dichlorolichexanthone (major), 2,4,5-trichloro-
lichexanthone (major), 2-chlorolichexanthone (minor), 2,4-dichlorolichexanthone (minor
to trace) and stictic acid (major).
Specimens examined:
AUSTRALIA, New South Wales, Minnie Water Beach, ca. 40 km ESE of Grafton,
29°47'S, 153° 18'E, alt. ca. 10 m, on Banksia behind beach, A.W.Archer P 382,
13.1x.1992 (NSW).
NEW ZEALAND, North Island, Three Kings Islands, Great Island, Castanay Valley, on
kanuka, D.J.Galloway s.n., Nov. 1970 (CHR 451403 pp).
Pertusaria subisidiosa is characterised by the isidiate thallus, the asci with four rough
spores, the conspicuous black ostoles and its chemistry. Pertusaria subisidiosa was
originally reported to contain stictic acid and traces of 4,5-dichlorolichexanthone (Archer
1991:242) but liquid chromatography showed the presence of the chloroxanthones listed
above. Pertusaria subisidiosa resembles the Northern Hemisphere species P. coronata
(Ach.) Th.Fr. but is chemically distinct from that species; P. coronata contains 4,5-
dichlorolichexanthone and stictic acid.
214
Pertusaria subplanaica Archer & Elix, Mycotaxon 45: 422 (1992).
Type: Australia, New South Wales, Patterson River [near junction with Hunter River,
ca. 10 km E of Maitland, 32°44'S, 151°33'E], J. L. Boorman s.n., Aug 1906; NSW-
holotype.
Thallus pale olive green somewhat cracked, surface smooth and dull, lacking isidia and
soredia, corticolous; apothecia verruciform, conspicuous, dispersed, sometimes
confluent, concolorous with the thallus, flattened-hemispherical, constricted at the base,
0.8-1.5 mm diam.; ostioles inconspicuous, brown, | per verruca; spores 8 per ascus,
irregularly uniseriate, smooth, ellipsoid becoming fusiform, (60-)80-100(-120) »m long,
30-40 ym wide.
Chemistry: K-, KC-, C-, Pd-; 4,5-dichlorolichexanthone and 2,2'-di-O-
methylstenosporic acid, with traces of planaic acid and 2,2'di-O-methyldivaricatic acid.
Specimens examined:
AUSTRALIA, Queensland, Mt. Mee State Forest, J. Hufellner 16887, 13.viii.1986
(GZU); Gambubal State Forest, E of Emu Mtn, J. Hafellner 16339, 7.1x.1986
(GZU).N.S.W.: Blue Mountains N.P., Mt. Wilson, K. Kalb 20468, 31.vii.1988 (herb.
KALB); Royal N.P., Bola Creek, K. Kalb 2/690, 21700, 2.viii. 1988 (herb. KALB).
NEW ZEALAND, North Island, Auckland City, Rangitoto Island, J. Wadham s.n.,
19.1.1951 (AK 172648); Thames County, B. Hayward s.n., Aug.1973 (AK154526);
Coromandel County, Mt. Mangatawhiri, 36°47'S, 175°46'E, B.Hayward s.n.,
Aug.1974 (AK171724).
Pertusaria subplanaica 1s characterised by asci with eight spores and the presence of
2,2'di-O-methylstenosporic acid. It occurs in eastern Australia and the North Island of
New Zealand.
Pertusaria thamnolica Archer, Mycotaxon 44:16 (1992)
Type: Australia, New South Wales, E side of Mooney Mooney Creek, ca. 8 km WSW of
Gosford, 33°26'S, 151° 15S'E, alt. ca. 10 m, on Casuarina, A.W. Archer P 178,
24.v.1991; NSW-holotype; ANUC-isotype.
Thallus off-white to pale grey, areolate and cracked, surface smooth and dull, lacking
isidia and soredia, corticolous; apothecia disciform, conspicuous,scattered, 0.5-1.5 mm
diam., with thick inrolled margins, surface of disc white-pruinose; spores 8 per ascus,
uniseriate, ellipsoid, smooth, 22-32 xm long, 12-17 ~m wide, spore wall single, ca. |
pm thick.
Chemistry: K+ yellow, KC-, C-, Pd+ yellow; thamnolic acid.
Specimens examined:
AUSTRALIA, New South Wales, Buckenbowra River Estuary, 7.5 km W of Batemans
Bay, 35°42'S, 151°06'E, on Casuarina, K.Kalb 18238, 1892], 4.viii.1988 (herb.
KALB); ibid., J.A. Elix 22360, 4.viii. 1988 (ANUC); Jerrawangla State Forest, Twelve
Mile Road, near Boyd Lookout, 35°10'S, 150°23’E, alt. ca 500 m, on Leptospermum,
A.W. Archer P 290, 29.ii1.1992 (NSW); Gibraltar Ranges National Park, track to the
Needles, 29°31'S, 151°22'E, alt. ca. 400 m, on fallen branch, A.W. Archer P 440,
28.viii. 1992 (NSW).
215
NEW ZEALAND, North Island, Egmont National Park, Ngatoro Track, 26 km S of
New Plymouth, 39°15'S, 174°07'E, alt. 590 m, on fallen branch, J.A. Elix 19232,
11.1.1985 (ANUC).
South Island, Lewis Pass area, Nina Valley terrace, 42°18'S, 172°23'E, alt. 700 m, W.
Malcolm 743, 1.i1.1993 (CHR 470254).
Pertusaria thamnolica resembles P. truncata Krempelh.but is distiguished from that
species by the presence of thamnolic acid and the slightly larger spores. The species is
known from New South Wales and the North and South Islands of New Zealand.
Pertusaria velata (Turn.) Nyl., Lichenes Scandinaviae: 179 (1861)
Parmelia velata Turner, Trans. Linn. Soc. London 9: 143 (1808).
Type: Great Britain, Sussex, on ash tree, W.Borrer s.n., 1805; BM-holotype
Thallus greyish-white to off white, slightly cracked and areolate, surface smooth to
slightly wrinkled, dull, lacking soredia and isidia; corticolous; apothecia disciform,
numerous, scattered or crowded, concolorous with the thallus, 0.5-1 mm diam., discs
pale to dark reddish-brown, slightly or densely white pruinose; spores | (rarely 2) per
ascus, ellipsoid, smooth, thin-walled, 100-185 ym long, 30-45 um wide.
Chemistry: K-, KC+ orange red, C+ red, Pd-; lecanonc acid
Specimens examined:
NEW ZEALAND, South Island, Nelson, Cobb River Dam, Mason s.n., Feb. 1959
(OTA 599a, 600a); Maruia Springs, W of Lewis Pass, alt. cu. 400 m, P. Child 1300,
6.1.1971 (CHR 452779); Lewis Pass, Nina valley, 42°28'S, 172°23'E, alt. ca. 700m,
Nothofagus dominated river terrace, on fallen branch, W. Malcolm 372, 2.11.1993 (CHR
470422); Mt. Arthur, Flora Hut road, 0.7 km W of Flora Saddle, 41°11'S, 172°44'E, alt.
ca. 950 m, W. Malcolm 872, 13.v.1993 (CHR 470256).
Pertusaria velata is a wide-spread species with a pantropical and temperate distribution
(Dibben 1980:78) characterised by disciform apothecia and the presence of lecanoric acid.
Although the asci usually have one spore, two-spored asci were also found in the New
Zealand specimens.
KEY TO CORTICOLOUS NEW ZEALAND SPECIES OF PERTUSARIA
| Detailed descriptions of species in the key that are not reported
above are given in Galloway (1985)|
1. Thallus lacking spores, with sterile discs oF SOralia...........cecceeeeeeseeeeeeeeeeeeeeee 2
Thallus with spores, in disciform or verruciform apothecia.............:ccsseeeneeeeeeD
2. Thallus sorediate, with white soralia, K-, Pd+ yellow, psoromic acid....P. psoromica
DBhallus with sterile paises ccc vette eeuts csc sesise eyes te ek a meee nas cee ce
baDisesuK- KC+ violet:: picrolicheniGgacid ss. cctv aavercyt. Gesrsttet-- P. truncata
Discs Rta violétvorsyellow gh Geaind 28 deucmetenayerrclry) sets ioe seein... «80: 4
4. Discs K+ violet, Pd-, hypothamnolic acid..........0........ss00seeeess P. novaezelandiae
Discs K+ yellow, Pd+ yellow, thammolic acid......... sc eseceeseeeeees P. thamnolica
Deen DOME Clam CISCIFOFM a. .c4..1 14s eer ee. erat em areas CueEE Ae ooh Se eee 6
ADOtNECLARVErrUCILOlN) tca.ssie essere eee AO, Cae I cc sn res 12
216
GEO DSPOTeS) 7S PPE NwasCuse AL Aves te me ae Meee eter ear tee es uke ae ee mee we om 7.
SPOres hf Per ASCUSE A ioaeesar esc tees Nee oe aed k skis SoON a Alar dc eh RMeN ENT Cale ane ete i 9
7. Spores spherical, 15-20 wm diam.; discs K-, KC-; lichen compounds absent..........
b Ket'edisls ialed esa bids sue ditate ra cines epaenee sUbk VNR adie ome tele Ree a meee Teena P. sporellula
Spores ellipsoid, 20-32 um long; discs K+ yellow or KC+ violet...........ccccceees 8
32) Dises* Ke, RCP widtet Pdsnisk oe ae cree rest ue ance ee P. truncata
Discs) K+ yellows | KG=:) Pde yellow uncer teen P. thamnolica
OL Diss) Regs Gani Ro BTR Sa a lia ea ae al rec en let ere ae er 10
Dises'(K+-" violeti or K+ yellow iin. citi ec escenes cor oder ccstenest uetonae ene 11
LOL CDSS) Ge suey Ua CHOP CA Ln Rnb heaters Sie nl wea er oe P. circumcincta
Dises?G+"red!"lecanoric acids Rae a eee ee P. velata
11. Dises K+: violet, Pd-; hypothamnolic ‘acid.......0 0c) cco. P. novaezelandiae
Discs K+ yellow, Pd+ yellow; thammolic acid; ....... cee eeeeeeeee P. scutellifera
I2Z°S pores 2h (or rarely 3)7 Perm asCus icc. dances terse ncaateer ice eee mene cae 13
SPOres 4" Or Sw peri ASCUs.s A aac sees aaeeel cuca ot ee eat eee RS Ee Rea 7
13k) Spores: S150) peme LONER A SS Oe RSs sky ee eee 14
SPOTS! cb DO WAV LON Gee oecctk sae seeseet etl cuss ec mee tevg ee ee eae eee ene 15
14. Verrucae conspicuous, raised; spore wall rough; spore length 160-250 pm...........
SA eolod geet g stone eettas Oe daet ease cn scr ctge dee nets UN aE tek neU te eer aie or P. sorodes
Verrucae inconspicuous, immersed; spore wall smooth; spore length 210-300 pm
EaOvaa SOMES Laer R ERs aah nah ya tee, deta tele ae dear e nth Cina acyl s aemmntn tte P. hadrospora
15. Ostioles pale, translucent; thiophaninic and stictic acids............... P. schizostomella
Ostioles black*stictie*acid absent.ca oiics ccs utecesc cot coeca tre eaane ee eee ene 16
16. Thiophaninic acid present; spores rarely 3 per aSCUS; ...........c.ccceeeees P. alboatra
No lichen compounds present; spores always 2 per ascus; ........... P. melaleucoides
17. (12) Spores 4 per ascus, spore walls rough; thallus isidiate.............. P. subisidiosa
Spores 8 (sometimes 6-8) per ascus, spore walls smooth; thallus not isidiate........ 18
18. Spores predominantly uniseriate, ostioles black or pale..............ccccccccessssreees 19
Spores: predominantly: biseriate; ostioles? DIACK 1.0.2. Lail 02. nuskssgaeecsaeesneeunne 23
19: Ostioles: black; ConmspiCuous.c2 UA. ibs A iat see ene uch cocereees een ar tem nae 20
Ostiolespaley WICONSPICUOUSA., c\h.tiemcesaecdensccds pede entagerte Aer eearn ee ee 21
20:Spores 50-70) pay tone thiophaninic V acid 0 oi2a.c..cctssesdeteee reser P. laevis
Spores 30-37 pm long; 4,5-dichlorolichexanthone.............ccceseeceeesees P. parvula
21. Thallus lacking lichen compounds; spores 50-62 pm long...........0000 P. albissima
Thallus with lichen compounds; spores 70-100(-120) pom long..........cceeeeees ae
2200220 -methyistenosporic-.acid present:/)..-2,-cet haicesasdnsctte enasevicetas P. leucoplaca
z2'\2-di-O-methyistenosporic acid? present... 2. .5..02.ccctewssesoce awe P. subplanaica
230418): Spores<sO\ gam “POM isl. d le. ces ods cedeadancmareesse reeds snae treme ees cameaes 24
Spores 4>SO0i per LOM ceca aad aes a raat cee cpt Penge tenet gauetnee 2D
217
24. Thallus KC+ yellow, arthothelin & thiophanic acid; ......... eee P. bartlettii
Thallus KC] slacking Jichen cOMmpOUNGS;. ie. ayecpeeessene nny ateuiexsneuen P. leucodes
Porpnalus K+ red, Pods yellow: NOrstictic ACIC Aaa cadereectnensavs goceseses P. celata
hhallus Ke Pd=s thiophaninic, acids\Uo. nie ec smerecta: serenei P. theochroa
ACKNOWLEDGEMENTS
The authors are grateful to the Herbaria referred to above and to Dr. W. Malcolm
(Nelson) for the loan of specimens, to Mr. D. Verdon for checking the Latin descriptions
and to Ms. C. Barclay (née Crook) for technical assistance with liquid chromatography ;
one of us (AWA) is grateful to the National Herbarium of New South Wales for
arranging the loan of type and other specimens and for permission to use the facilities of
the Herbarium; the other (JAE) gratefully acknowledges the Australian Research Council
for generous financial support.
LITERATURE CITED
Archer, A. W. (1991). New species and new reports of Pertusaria (Lichenised
Ascomycotina) from Australia and New Zealand. Mycotaxon 41, 223-269.
Archer, A.W. & Elix, J.A. (1993). Saxicolous species in the genus Pertusaria (Lichenes)
common to New Zealand and Australia. V.Z.J. Bort. 31, 111-116.
Archer, A. W. & Elix, J. A. (1993a). Additional new taxa and a new report of Pertusaria
(Lichenised Ascomycotina) from Australia. Mycotaxon 49, 143-150.
Dibben, M. J. (1980). 'The chemosystematics of the lichen genus Pertusaria in North
America north of Mexico’. Milwaukee Publications in Biology and Geology,
Number 5 (Milwaukee).
Elix, J.A & Venables, D.A. (1993). 4-O-Methyllividic acid, a new lichen compound.
Mycotaxon 47, 275-281.
Galloway, D.J. (1985). 'Flora of New Zealand Lichens', Government Printer,
Wellington.
Krempelhuber, A. (1876). Neue Beitrage zur Flechten-Flora Neu Seelands.Verhandl.
cool.-bot. Ges. Wien 26, 447-460.
Miiller, J. (1884). Lichenologische Beitrage XIX. Flora, 67, 299-306; 349-354.
Nylander, W. (1888). Lichenes Novae Zelandiae. P. Schmidt, Paris.
Oshio, M. (1968). Taxonomical studies on the family Pertusariaceae of Japan. J. Sci.
Hiroshima Univ. Series B, Div. 2, 12, 81-163.
Vainio, E.A. (1907). Lichenes novi rarioresque. Ser. 1V. Hedwigia 46: 169.
Zahlbruckner, A. (1941). Lichenes Novae Zelandiae a cl. H. H. Allan eiusque
collaboratoribus lecti. Denkschr. Akad. Wiss. Wien math.-naturwiss. K1.104,
249-380.
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Volume L, pp. 219-270 January-March 1994
ALTERNARIA THEMES AND VARIATIONS (74-105)
EMORY G. SIMMONS
717 Thornwood Road, Crawfordsville, IN 47933
ABSTRACT
This paper includes clarification of an earlier paper
on euphorbiicolous Alternaria species (Simmons 1986), most
notably on Euphorbia pulcherrima and E. marginata, and
expansion to include opinions on all taxa of Alternaria and
somewhat similar genera that have been described from
euphorbiaceous substrates. Proposals of A. acalyphae (Nelen)
Simmons, comb. nov. (= Macrosporium acalyphae Nelen; on
Acalypha australis); A. obtecta Simmons, sp. nov. (on
Euphorbia supina and E.marginata); A. constricta Simmons, sp.
nov. (on EE. marginata); A. agripestis Simmons & Mortensen,
sp. nov. (on E. esula); A. subulata Simmons, sp. nov. (on E.
hirta); A. acalyphicola Simmons, sp. nov. (on Acalypha
indica); A. heveae Simmons, sp. nov. (on Hevea brasiliensis) ;
A. cicina Simmons, sp. nov. (on Ricinus communis).
XIII. (MOSTLY) ALTERNARIA ON EUPHORBIACEAE
Introduction
Taxonomic interest in euphorbiicolous species of Alter-
naria in the past decade has included incitants of a disease
affecting commercial production of poinsettia (Euphorbia
pulcherrima Willd. ex Klotzsch) and of a disease of an agro-
nomic pest, leafy spurge (E. esula L.). Two papers published
within two months of each other in the winter of 1985-1986
(Yoshimura et al. 1985; Simmons 1986) discussed Alternaria
taxa on species of Euphorbia. Both papers contained propo-
sals for combining the epithet of Macrosporium euphorbiae
Bartholomew (1908) with Alternaria, and each selected a
lectotype specimen for the name from among specimens distrib-
220
uted by Elam Bartholomew in Fungi Columbiani No. 2633 (1908).
The proximate dates of publication (December 1985 and
February 1986), differences in author interpretation of Bar-
tholomew’s species because of reliance on different portions
of his original collection(s), and the misapplication of the
name A. euphorbiae to the poinsettia pathogen (correct name
= A. euphorbiicola) are creating needless confusion in perti-
nent herbaria, reference culture collections, and derivative
publications (e.g., Holliday 1992, and abstracting journals
such as the Review of Plant Pathology).
The following observations are organized to present:
1. Opinions on the identity and nomenclatural status
of each taxon described in Alternaria, Embellisia, Macrospo-
rium, Stemphylium, and Ulocladium (plus Botryosporium
prorumpens Schweinitz) whose substrate is euphorbiaceous.
2. A more detailed discussion of the typification of
A. euphorbiae (Barth.) Aragaki & Uchida, particularly with
reference to the hazards of studying different packets of an
exsiccata issued in 70 sets of supposedly duplicate or
multiple units, as Bartholomew is reported to have done for
Fungi Columbiani (Stevenson 1971).
3. The results of a recent review of approximately 130
fungus specimens and isolates from euphorbiaceous substrates,
primarily from BPI and IMI, but also from NY and other
institutional herbaria as well as from several individual
contributors.
The number of novelties described below will not seem
extravagant when it is realized that conidia of at least four
taxa of Alternaria are present on the collection of spotted
leaves of Euphorbia marginata Pursh chosen by Aragaki and
Uchida as lectotype of A. euphorbiae, and that the single
genus Euphorbia, with its 2000 species, contributes less than
half of the taxa suggested for the Euphorbiaceae, with its
300 genera and 5000 species (Willis 1985).
Conventions used in this discussion include: herbarium
and culture collection abbreviations are from Holmgren et al.
(1990), given here in boldface type; references to my records
of material studied are in record volume-page format, e.g.,
EGS 41-034; ICBN refers to the International Code of Botani-
cal Nomenclature (Greuter et al. 1988, except when an earlier
221
edition is specified); an abbreviation "Alt.T&V" often is
used to simplify reference to this series of papers on
Alternaria (Simmons 1981-1993), usually with a number speci-
fying a pertinent unit in the series; routine culture condi-
tions include PCA, Hay, and V-8 agars (Simmons and Roberts
1993, p. 136) in plastic Y-plates at ca. 22C under a 10/14hr,
cool-white fluorescent light/dark cycle; microscopic details
are recorded from observations made at 195X and 780X.
Chronological Overview of Taxon Names Attributed to
Phaeodictyosporous Hyphomycetes Described on Euphorbiaceae
74. 1832 Botryosporium prorumpens Schweinitz
Trans. Amer. Phil. Soc., n.s., 4: 306. In Euphorbiae
epidermide...Bethlehem [Pennsylvania].
Lectotype: FH, Botryosporium prorumpens, Schw! in
Euphorbia & Diantho; Herb. Schw. (EGS 38-026).
The original Schweinitz specimen associated with this
name bears several different fungi. The collection itself
has been subdivided over the years, with portions now held
at least in BPI (herb. Michener); FH (herb. Curtis); K (herb.
Berkeley); and PH (herb. Schweinitz).
Damon (1952) reported that a fungus in the BPI portion
appeared to be “of the general type found in Stemphylium" and
illustrated two spores that could belong to one of several
different genera but not to Stemphylium Wallroth. A few
very scarce conidia found on the BPI portion (EGS 17-117) are
of an Alternaria of the A. alternata group.
The FH portion (EGS 38-026) bears some rare alternari-
oid elements similar to those of the PH portion (see below),
some conspicuous clumps of miscellaneous fungus’ spore
skeletons (presumably deposited by insects, and illustrated
in a sketch on the herb. Curtis specimen label), and a moder-
ately extensive group of prorumpent sporodochia of Epicoccum
(conidia also sketched on the label). I have not examined
the K portion, but the records at the International Mycologi-
cal Institute (IMI) indicate that M. B. Ellis at some time
had given the opinion that the fungus on the K portion is an
Epicoccum.
The PH portion (EGS 12-085) bears numerous phaeodictyo-
222
conidia lying detached on the substrate. These are striking-
ly similar to Alternaria conidia, but there is no readily
apparent pore or pigmented annulus in the broadly rounded end
of the spores; the narrowly tapered end of each of the few
conidia examined is fractured. The visible evidence is
inadequate to support calling the fungus an Alternaria,
although it may be.
Schweinitz (1832) described a fungus with rudimentary
sporodochia arranged in two to several subconfluent series
and bearing very dark fascicles of dark, compressed-globose,
pedicellate spores (which he illustrated as being dictyo-
spores). His description and illustration are consistent
with the characters of Epicoccum Link, although the conidia
in his drawing could be interpreted as inverted alternarioid
entities. The FH portion of the original specimen bears a
good representative of Epicoccum. This FH portion of the
original collection is chosen here to be the lectotype of
Botryosporium prorumpens Schw.
Monotypic Botryosporium Schw. (1832) is a later homonym
of Botryosporium Corda (1831), whose own type, B. diffusum
(Albertini & Schw.) Corda, probably is conspecific with B.
longibrachiatum (Oudem.) Maire (fide Mason 1928). The genus
name Botryosporium Schw. is illegitimate (ICBN Art. 64); its
type is a species of Epicoccum.
ioe 1878 Macrosporium compactum Cooke
Journ wiiLinn! wsoct" “Botany yl] AZ, On stems of
Ricinus: “Houston (Rav 2272); (2/3.)%:
Lectotype: BPI 0445206, Macrosporium compactum Cke.,
on. Ricinus) “Houston, Texas, Apy.;' 16," U869." HoWen 92757 ne
(EGS 07-109).
Cooke (1878) described this fungus as dark and effuse
on a compact crust, with simple, brown, septate, fasciculate
conidiophores, and with oval, obtuse, sooty brown, cellular
conidia 20-30 x 12-14 um.
Cooke also illustrated his concept of M. compactum in
a plate of figures accompanying a paper on New Jersey fungi
(Cooker ands BL Iisi( L878 Pi 99k). toe Loy Although this
illustration cannot be tied directly to the typification of
M. compactum, the proximate dates of the two publications
should give weight to Cooke’s linking a description and an
23
illustration so immediately with a single species name.
The Ravenel specimen bears an intimate heterogeny of
sporulation elements of at least Alternaria, Curvularia,
Pithomyces, and Ulocladium taxa. Cooke's description of
sooty brown (fuligineis) conidia with obtuse ends and, in
particular, his illustration suggest identity with the
Pithomyces that is present. Certainly the rather pale,
sharp-beaked Alternaria conidia that are present in abundance
do not fit the description.
McClellan (1944), ignoring Cooke's specific character-
ization of shape, pigmentation and size of conidia, selected
conidia from "herbarium type material" (unspecified) that
approximated the spore characters of an Alternaria isolate
from Ricinus leaves that he was studying. He proceeded to
propose the new combination Alternaria compacta (Cooke)
McClellan (see Alt.T&V 85 below).
The lectotype of M. compactum Cooke, chosen here from
among specimens listed in the protologue, is a species of
Pithomyces Berk. & Broome not readily if at all distinguish-
able. from P. chartarum (Berk. & Curt.) (M2 B.. Ellis on’ the
basis of the conidia seen.
Vor 1886 Macrosporium trichellum (Arc. & Sacc.) Arc. &
Sacc. var. caulicola Saccardo [as "caulicolum" ]
Sylloge Fungorum 4: 525. In caulibus emortuis Euphor-
biae biglandulosae, La Mortola, Ital. bor. [Penzig].
Type specimen not found. Variety typified by the
Saccardo description, which he copied from observations made
by Penzig (1884) on a Penzig specimen discussed in his Funghi
della Mortola.
Arcangeli and Saccardo (1878) described Stemphylium
trichellum Arc. & Sacc. on the printed label of a specimen
distributed as no. 698 in Series II of the exsiccata Erbario
Crittogamico Italiano (1858-1885, various authors and pub-
lishers). They observed that the fungus, found on leaves of
Euonymus japonicus L., had 3-5 septate-muriform, fuligenous,
ellipsoid to obovoid conidia 30-35 x 18 wm, that it agreed
in some respects with Stemphylium pyriforme (Corda) Bonorden
and a few other dark-spored taxa, and that they wondered if
Stemphylium, Macrosporium and several other similar genera
(named) really differed essentially from each other.
224
Later Saccardo (1881) issued an illustration of this
fungus in his Fungi italici autograph. del. no. 853 but under
the name "Macrosporium trichellum Arc. et Sacc.," citing its
earlier publication as a Stemphylium. The illustration and
conidium dimensions match the original 1878 description.
Penzig (1884), in his Funghi della Mortola, identified
one of his specimens collected on stems of Euphorbia biglan-
dulosa as M. trichellum Arc. & Sacc. and gave dimensions of
conidiophores as 35-50 wm long, of conidia 33-35 x 16-18 um.
Finally, Saccardo (1886) considered Penzig’s fungus to
be a distinct variety of the original species because of its
comparatively short conidiophores and caulicolous habit on
a host different than that of the original taxon. He created
the name M. trichellum var. caulicola Sacc., though there is
no evidence either in his entry in the Sylloge Fungorum or
in his herbarium at PAD that he examined Penzig’s specimen.
I have not been able to locate Penzig’s collection of
this fungus. Without it, it is not profitable to speculate
on the nature and identity of the fungus he collected, even
at the genus level.
Ws, 1887 Macrosporium commune Rabenh. f. Ricinis communis
in Berlese and Roumeguere: Revue Mycologique 9: 180.
The entire entry reads "4240 (38) Macrosporium commune
Rabh. f£f. Ricinis communis. In foliis emortuis. Cerra de S.
Bento pr. Coimbra: Juan. 1887."
The name of the forma, if such was the intention, is
not accompanied by any sort of diagnosis or description. It
is a nomen nudum. The numbers "4240 (38)" refer to specimens
issued in Roumeguere's Fungi selecti exsiccati. The unit of
this exsiccatus that I have examined (NY; EGS 05-075) bears
hyphal fragments but, in the single slide preparation
examined, no conidia pertinent to this discussion. Addition-
ally, the name, as a forma, would have no priorible status
in nomenclature.
Se 1908 Macrosporium euphorbiae Bartholomew Figewl
E. Bartholomew, Fungi Columbiani no. 2633, with type
description on printed label of exsiccata packet. On
living & languishing leaves of Euphorbia marginata.
Fig. 71. Alternaria euphorbiae:
Macrosporium euphorbiae Barth.
(BPI 0445288).
conidia from lectotype of
Bar = 50 wm.
226
Lectotype: BPI, certain conidia (Fig. 71) on slide
#1 of M. Aragaki and J. Y. Uchida, maintained in a separate
packet associated with BPI 0445288, a unit of Bartholomew's
F. Col. 2633, and with an annotation slip "#1 This specimen
designated as lectotype of Alternaria euphorbiae (Barth.)
Aragaki & Uchida. Published by M. A. Yoshimura, J. Uchida
& M. Aragaki in Plant Disease, March, 1986 70: 75." (EGS 41-
116). [The cited paper appeared in Plant Disease 70 (January
1986): 73-75, which actually was issued in December 1985. ]
The herbarium of the National Fungus Collections (BPI)
contains three packets/units of F. Col. 2633. Aragaki and
Uchida apparently indicated their lectotype choice to BPI but
in their published treatment did not specify the actual
specimen or the institution involved, stating "The lectotype
specimen (designated by M. Aragaki and J. Y. Uchida) fit the
description given by Bartholomew, . . ." [The Interna-
tional Code of Botanical Nomenclature in effect in 1985-1986
(Voss et al. 1983) did not require specific mention of the
herbarium or institution in which a designated lectotype is
conserved. The edition of the ICBN published in 1988
(Greuter et al.) does make such a requirement in Art. 8.4,
which became effective 1 January 1990. ]
Simmons and Francis (Alt.T&V 14, Simmons 1986) also
published observations on Macrosporium euphorbiae Bartholomew
and, unaware of the Aragaki and Uchida work, selected as
lectotype a fungus found in a different unit of F. Col. 2633
(NY; EGS 03-047). Regardless of the merits or differences
of observations made by the two different pairs of authors,
the Aragaki and Uchida prior choice of lectotype must
prevail, and fungus elements present in BPI 0445288 must
serve as lectotype of M. euphorbiae Barth.
However, definition of M. euphorbiae within the context
of this a.cepted lectotype choice raises problems for the
simple fact that the slide preparation #1, representing the
material examined microscopically as a basis for the lecto-
typification, actually bears conidia of three different
species of Alternaria and some large, long-beaked conidia of
a Nimbya taxon. The nature of the three distinguishable
kinds of Alternaria conidia is taken up in detail in a
separate section following this chronological treatment of
names (Alt.T&V 97-99). Observations in that place support
a caution that all mycotaxonomists (including me) need to
heed when dealing with exsiccatae of microscopic fungi in
particular, viz., that a fungus name refers to a fungus of
227
a certain circumscription, not to a host lesion of a certain
nature, even less to detached portions of field material in
a common packet, and most certainly not indiscriminantly to
all the putative “isotypes" or "duplicates" distributed by
the score in major exsiccatae, only a few of which the
distributor could possibly have examined in detail.
The main point in this chronology is to define M. eu-
phorbiae as critically as possible on the basis of Barthol-
omew’s type description and the lectotype source. It is not
essential at this point to discuss the characters of the
Alternaria pathogen of poinsettia discussed in the Yoshimura
et al. paper (1985) and the Simmons paper (1986) or whether
it is the same taxon as M. euphorbiae Barth. (although it is
not).
Bartholomew described Macrosporium euphorbiae in as
much detail as was customary for the time. The appearance
of the leafspots of the host are not diagnostically helpful,
as at least one such lesion provided conidia of the three
alternarioid fungi of the lectotype slide. The spots were
described as, and are, "amphigenous, brown, suborbicular,
subcircinate, lighter at the margin," and in this lectotype
unit ca. 3-8 mm diam.
The original description of conidiophores likewise is
not specifically applicable because it is too generalized to
permit application to one of the conidial types present but
not the others. Conidiophores ("hyphae") were given as
"amphigenous, dark brown at base, lighter toward the apex,
dense, fasciculate, simple, divergent, 2-6 septate," which
some are, some are not. |
The description of conidia thus becomes the only
Significant portion of the characterization that permits a
choice of the M. euphorbiae lectotype from among the diversi-
ty of taxa present. Bartholomew continues: "Conidia light
brown, obclavate or subelliptical, 4-8 septate, 40-75 x 12-
2Qz, with frequent longitudinal septa. Apical appendages
flexuous, curved, as long or longer than the conidia."
Figure 72 illustrates four kinds of conidia present on
the lectotype slide that are pertinent to this discussion.
The three Alternaria taxa are treated in more detail in
VOWS OR AN OR SS oe ES Be de
228
Fig. 72a An Alternaria species with fully developed
conidium body 60-80 x 15-18 yum, with 7-9 transverse septa and
one, rarely two longitudinal septa in 2-5 of the largest
transverse segments of the spore. The outer wall appears
smooth and the transverse septa appear distinctly medium
brown in color. Apical narrowly filamentous beaks are almost
straight to smoothly flexuous, 110-170 x ca. 2 um.
These conidia represent typical M. euphorbiae Barth. ,
i.e., Alternaria euphorbiae (Barth.) Aragaki & Uchida, in
that they fit the type description well, and certainly better
than other conidia present on and illustrated from the
lectotype slide. If it should be argued (which I do not)
that Bartholomew may have seen and included the other kinds
of conidia in his species concept, and that one of them
should be designated lectotype, then it is specified here
that only the kind of conidia in slide #1 associated with BPI
0445288, as characterized above for Fig. 72A and illustrated
here in Fig. 71, are designated as lectotype of M. euphorbiae
Barth. (See ICBN, Art. 8.1(b), Greuter et al., 1988.)
Rig 272b An Alternaria species with fully developed
conidium body 70-85 x 18-20 wm, with 9-12 pale, indistinct
transverse septa and one obscure longitudinal septum in 0-2
of the transverse segments. The outer wall is pale, almost
hyaline, as are the septa, which seem to be masked by a
crowded, extremely minute, punctulate ornamentation. Apical,
narrowly filamentous beaks are straight or somewhat curved,
LTO- 120"x ican 2m,
The conidia of this species, with their large body and
filamentous beak, are distinguishable additionally in
pigment, pattern and indistinctness of septation, and wall
ornamentation from those of M. euphorbiae.
This taxon is known and has been isolated from other
euphorbiicolous sources. It is discussed in more detail
below in Alt.T&V 98 as a new species.
Fig. 72c An Alternaria species with fully developed
conidium body 35-50 x 10-12 ym, with 6-7 transverse septa and
usually 1-2 longitudinal septa in each transverse segment
except the apical and basal. Wall pigmentation is medium
yellow-brown; walls are punctulate roughened overall.
Conidia are ellipsoid, tapering apically into a beak cell ca.
18 zm long, or beakless with an apical secondary conidiophore
ca. 15 x 4ym, which characterizes the taxon as chain-forming
229
Fig. 72. Conidia of taxa on lectotype slide of Macrosporium
euphorbiae Barth. (BPI 0445288). a. Alternaria euphorbiae,
b. Alternaria obtecta, Cop Altermnarian sp.,4 ~d.eNimbyacsp.
Bar = 50 pm.
230
in nature.
This conidial component of the lectotype slide has only
generic resemblance to conidia discussed in Fig. 72a and b
above. It is readily separable from the others in the
context of this slide, but there are insufficient numbers of
conidia for speculation at present as to its identity among
similar small-spored species.
Fig.) 72d A Nimbya species with fully developed conidium
body 50-70 x 13-18 ym, with 8-10 transverse, variously angled
lacunae partially or completely separated by transverse septa
and one longitudinal septum or lacuna partition in 1-3 of the
transverse segments. Wall color is a pale yellowish brown.
Apical filamentous beaks are straight to smoothly flexuous,
L00-170ux ‘cas /2) 2m.
The strongly lacunate, rarely longiseptate conidia of
this Nimbya contrast well with the typically muriformly
septate M. euphorbiae conidia in this slide preparation.
Here again, there are insufficient numbers of conidia for
comparison with other known species of Nimbya.
79. 1921 Macrosporium cavarae Parisi
Bulli Orto "Bot, WUniv, Napoli 650291) Inifotins Rrer:
communis.
Type: published description (l.c.); specimen lacking.
The fungus was described (in Latin) as: conidiophores
simple, flexuous, sooty brown; conidia clavate [obclavate?],
muriformly septate, with 4-8 transverse septa, somewhat
constricted, 34-40 x 10-13 uw, or 39-47 yw with pedicel
[beak?], concolorous [with conidiophores?]. Given the widely
held concept of Macrosporium in 1921, the description may be
of a relatively small-spored Alternaria with either a short
beak cell or a short apical secondary conidiophore 7-8 yum
long. In the absence of a specimen this is mostly specula-
tive, even as to genus. The taxon is unknowable on the basis
of the information available.
McClellan (1944), followed by Joly (1964), considered
M. cavarae identifiable with M. compactum Cooke, i.e., with
A. compacta (Cooke) McClellan. The type of M. compactum,
however, is a Pithomyces, which applies automatically to the
identity of A. compacta. (See Alt.T&V 75 above.)
231
80. 1921 Macrosporium euphorbiae Reichert
Bot. Jahrb. soyst:. \ 5677 2345) Hap deine fol iis, Euphorbiae
prunifoliae (Jack.) J. Muller prope Salamum prope
Mansurahm, 6 Decembri 1911 (leg. Schweinfurth in Mus.
DOE WBerol. ).
Holotype: B, “auf Euph. prunifolia, Salamum prope
Mansurah,, 6.12.1911. °S... Schweinfurth,." | (EGS 09-103).
The pertinent fungus present on leafspots of this
collection is an Alternaria. It was discussed and illus-
trated in Alt.T&V 15 as A. euphorbiicola Simmons & Engelhard
(Simmons 1986), a new name for M. euphorbiae Reichert, which
is a later homonym of M. euphorbiae Barth. (1908).
Largest conidia (ca. 50-60 x 18-20 um) of the Reichert
type specimen are beakless or have a short tapering beak cell
up to ca. 10um long. Many conidia of the type have produced
an apical secondary conidiophore 3-30 x ca. 3 wm. These
conidiophores are in no respect the equivalent of the long,
sinuous, filamentous beaks seen in the holotype of Bartholo-
mew's M. euphorbiae. The sturdy, stiff pseudorostra charac-
terize M. euphorbiae Reichert as chain-forming in nature.
The fungus described by Reichert is known from addi-
tional herbarium specimens (e.g., from Hong Kong, Tanganyika,
U.S.A.) as well as from isolates originating in Florida and
Hawaii. This material, observations on it in culture and its
relationship to a disease of poinsettia are discussed in
Alt.T&V 15 (Simmons 1986) and 95 below under its correct
name, A. euphorbiicola.
81. 1929 Macrosporium ricini Yoshii 1 nO) 6
BubU eeh. (RakteyTerkule: i Kjusu’ Impi Univ) 3e0'330:
[Engl. Summary 3: 330.] Occurring on the leaves of
castor oil plant (Ricinus communis L.), collected at
Suigen, Chosen, August 1926, September, 1927, at
Hakozaki, Fukuoka, July 16, 1928 (type), September,
1928.
Lectotype: BPI 0445446, [collected on] Ricinus
communis [at] Fukuoka [on] July 1928 (EGS 41-106).
I have had the opportunity to examine two Yoshii
specimens, the one deposited at BPI and one of later date,
wihy 516, 19295 lent to me.by),. Yoshii in 1956. with the label
Doz
"Macrosporium (Alt.) ricini Yoshii/Ricinus communis L. /Hako-
zaki, Fukuoka, H. Yoshii" (EGS 08-164). The date of the BPI
specimen is within the protologue data for the name and,
therefore, is chosen as lectotype. The label of the specimen
lent in 1956, although marked "Type," bears a date emendation
from August 22, 1930, to July 16, 1929, ‘which precludes
status as type when the date in the original description
clearly is given as month 7 of 1928 (Japanese text) and as
July 16, 1928 (English summary). The same fungus taxon is
present on leaves of the two different collections, but it
is necessary to specify the BPI material as lectotype in
accord with ICBN Art. 8 rules on lectotypification.
Yoshii described the fungus as incitant of a leafspot
disease characterized by irregular cinnamon-buff spots 10-20
mm. diam. that often are surrounded by a paler halo. His
description of the fungus fits [with ES comments noted] the
material in the specimen: fasciculate conidiophores light
brown to olivaceous, stout, 30-60 x 5.5-7.0 wm, 1-4 celled;
conidia greenish brown to olivaceous, fusiform to obclavate
with many [4-8] transverse and longitudinal septa, 42-78 x
8.7-19.5 wm, scarcely [nearly always slightly] constricted
at the septa, with slender hyaline beak 30-100 x 1.2-1.8 yum.
Yoshii’s work, reported in this 1929 paper, included
isolation of the fungus and inoculation of test plants, with
positive results. He was one prominent mycologist of the
time who noted correctly the relative positions of Alternaria
conidia and their conidiophores, viz., that the conidia are
obclavate with rounded base proximal to the conidiophore apex
and that the conidium’s apical extension is a beak, not a
pedicel at the base of the conidium. His excellent photo-
graphs (l.c., Tab. 2) leave no doubt as to conidium orienta-
tion and morphology.
B2.. 1939 Macrosporium rosarium Penzig var. piscariae
Sprague, in Sprague and Cooke, Mycologia 31: 51. On
living leaves of Piscaria setigera (Benth.) Peper,
South (of Many. (s') River, (Corvallis) wBenton Cow Ones
OSS ICA RLOVILO. Hives Jackson Sept, (2019145
Type: illustration and description, Mycologia 31: 49
(Fier n2 AS OC) apo. 1939. Two portions of the original
specimen have been examined in detail, BPI 0455448 (EGS 07-
1217 415108) srand@0SC) 1.091 618( ECS 12-08808 Not a single
conidium or conidiophore having the characteristics of the
200
Pete) Ol. Alternariva jriCini: conidia from lectotype of
Macrosporium ricini Yoshii (BPI 0445446). Bar = 50 un.
234
type figure and description has been found, even though
presumptive conidia and conidiophores of the kind of fungus
described (a Stemphylium) usually can be seen readily at 50-
60X dissecting microscope magnification. Sprague (in litt.
5-25-58) once suggested that a third portion may have been
sent to another herbarium. I have not encountered such in
searches for type material.
An excellent Alternaria and a Heterosporium are present
in abundance in the collection. Collector H. S. Jackson de-
scribed the conidia as "clavatis" in his manuscript notes,
so presumably he noted the Alternaria. However, the type
description, illustration, and reference to Macrosporium
rosarium Penzig (= Stemphylium rosarium (Penzig) E. Simmons
1990) undoubtedly suggest a Stemphylium. The type descrip-
tion indicates conidiophores 50-105 x 6-10.5ym with a series
of successive, inflated, internally proliferating conidiogen-
ous cells; pyriform to urn-shaped or subcylindric conidia
with muriform (usually 3 strong transverse septa), somewhat
constricting septation, are olive-brown, minutely and thickly
echinulate, 15-40 x 15-23 um.
Lacking a specimen and given the current inadequacy of
the systematic literature on Stemphylium, we now are unlikely
to be able to identify this taxon.
83. 1943 Alternaria ricini (Yoshii) Hansford Fig. 74
Proc. ‘Linn. Soo; London )153: 53.
Basionym: Macrosporium ricini Yoshii (1929). See
Alt.T&V 81 above.
Hansford offered the new combination in Alternaria on
the basis of Yoshii’s description and of a specimen (Hansford
8561) collected on Ricinus communis in Kampala, Uganda. He
added no novel information for the taxon except that "the
conidia are occasionally 2-catenulate." I have not seen the
Hansford specimen. It is worth noting that Yoshii’s type
specimen also contains a rare conidium with an apical scar,
indicating occasional production of short chains in nature.
Conidia in authentic Yoshii material as well as in
field specimens I have seen from several different countries
yield a microscopic picture suitable for correct determina-
tion of the species on natural lesions, but they are inade-
quate for identification of the taxon in axenic culture. The
Fig. 749) (Alternariairicini 3)\3i/conidiavat left) from original
field specimen; 6 conidia at right (all conspicuously orna-
mentea) trom culture 7d../on PCA. se Bari=)50yzm. Wi (EGS 06-181)
236
problem is not subtle (Fig. 74). Mature conidia from the
most abundant natural spore populations I have seen (EGS 06-
181 from Virginia, U.S.A.; and BPI 044651 ex IMI 68960 from
Southern Rhodesia (EGS 41-098)) tend to have a nearly
hemispheric or short, rounded obconoid base cell or region
and appear quite smooth at 780X. Conidia of comparable
maturity in cultures of EGS 06-181 on PCA and V-8 tend to
have a strikingly elongate base cell and a densely ornamented
outer wall. It is only on relatively non-nutritive Hay agar
that conidia similar in proportions to those from nature are
produced, and even these Hay-culture conidia are conspicuous-
ly ornamented.
The abundant sporulation of field specimen EGS 06-181
has conidia that are already narrowly appiculate when they
have only (3-)5 transverse septa (none longitudinal) and are
ca. 50 x 10 wm in size. The spore broadens and usually
lengthens to become 50-75 x 20-25ym, narrowing abruptly into
a fine, almost rigid beak 60-100 x ca. 1.5 wm. Largest
conidia are broadly ovoid to ellipsoid, with 8-9 transverse
septa and 1-2 longitudinal septa in each of most of the
transverse compartments. Conidium color is a dilute yellow-
ish tan; spore walls appear smooth.
An isolate derived from EGS 06-181 sporulates abun-
dantly on PCA and V-8 agars, less so on Hay. Conidia
produced on Hay agar within 1 wk. are, in general, similar
to those from the field specimen except that the conidium
wall is visibly ornamented. Fully developed conidia are
ellipsoid to long ovoid and ca. 65 x 18 wm with a shorter,
narrow beak. Secondary sporulation (chain formation) is rare
on Hay.
Conidia of EGS 06-181 produced on PCA (my usual compar-
ison medium) and V-8 agars differ strikingly from those on
Hay due to the presence of low excrescences evenly spaced,
almost crowded, over nearly all of the spore wall, even of
the smallest juvenile conidia. Spore bodies of most of the
fully developed conidia are long ellipsoid (widest in the
middle) or long ovoid (widest below the middle), with a
narrow beak. However, conidia with a terminal secondary
conidiophore (chain formers) commonly have an unusual obovoid
appearance, with the greatest diameter above the middle of
the spore body. Young narrow, transeptate conidia tend to
broaden variously as they mature without much change in the
diameter of the basal one or two cells. These cultural
characters of attentuate base and strong ornamentation of
Zor
conidia need to be added to the A. ricini species concept.
84. 1943 Macrosporium ricini Sawada
Rept. Agr. Res. Inst. Formosa no. 85: 97. Collected
as a parasite on Ricinus communis, Taihoku, Feb. 4,
1926, by Kaneyoshi Sawada [orig. in Japanese].
Holotype: TAI, on Ricinus communis L., Taipei (Tai-
DOK) erOr. wiheb.074. 5 9) 26% K. Sawada [modern label in
English] (EGS 12-101).
Sawada described a fungus with obclavate conidia
extending into a long beak, 3-11 transverse and 0-6 longitu-
dinal septa, 36-130 x 10-23 wm.
A single leaf with a distinct necrotic lesion well
darkened by conidia and conidiophores has been examined from
the type specimen. Most of the largest conidia have unusual-
ly swollen cells. No large conidium in this slide prepara-
tion retains an intact beak. The overall impression is of
A. ricini (Yoshii) Hansford in an overmature, overdeveloped
state.
Accepting M. ricini Sawada as identifiable with A.
ricini is a convenient and, I think, correct solution to the
handling of the Sawada taxon, for its name is illegitimate,
both as a later homonym of M. ricini Yoshii and in lacking
a Latin description or diagnosis (required by ICBN rules
beginning 1 January 1935).
65.) 1944 Alternaria compacta (Cooke) McClellan
Phytopathology 34: 229.
Basionym: Macrosporium compactum Cooke (1878). The
lectotype of the basionym is a Pithomyces, probably P. char-
tarum. See Alt.T&V 75.
86. 1959 Alternaria ricini (Sawada) Sawada
spec ss Frubla, Coll wAgr. Nat. slaiwan Univ.) no. 5 :, 208.
Basionym [illegit.]: Macrosporium ricini Sawada
(1943). See Alt.T&V 84.
The Sawada basionym gains no legitimacy by being
238
transferred to Alternaria, where A. ricini (Yoshii) Hansford
was established legitimately in 1943. The fungus collected
and described by Sawada appears to be an overdeveloped growth
of A. ricini (Yoshii) Hansford, but the names he published
illegitimately have no standing in mycological nomenclature,
not even as synonyms of the Yoshii/Hansford combination.
87. 1962 Macrosporium acalyphae Nelen Big ovo
Bot. Mater... OtdiiSporov.) Rast; Both inst. »Akad 7 iNauk
SSSR 15: 142. Habitatio. In foliis Acalyphae austra-
lis L. districtus Kalininensis pag. Rozhdestvenskoje
27 Viti L952 (typus)..
Holotype: LE? I have not seen the specimen designated
by Nelen as type and said by him to be conserved in LE. A
later Nelen collection was received from LE in response to
my request for Nelen types.
Most of the Nelen specimens lent to me by LE correspond
to respective protologues; a few do not. Having no reason
to believe that the type specimen of M. acalyphae would not
have been sent if it is extant at LE, it is reasonable to
conclude that it is not and that another specimen authentic
with Nelen and matching the type description can be chosen
as neotype of the name.
Neotype: LE, Macrosporium acalyphae Nelen, on Acalypha
australis, Amurskaya oblast, sepo Taibovka, pole [Amur
Oblast, Taibovka village, field], 1 IX 1959, E. S. Nelen.
(EGS 20-166).
Nelen described and illustrated a leafspot fungus with
oblong-oval, pale brown conidia with 9-12 transverse and 1-2
longitudinal septa, and an apical, hyaline, filiform beak
100-127 wm long. Dimensions of conidium bodies plus beaks
are 160-220 x 13520 zm. Nelen illustrated but did not
mention that the conidium beak may be 1-branched and may have
up to 8 septa more or less evenly spaced in the narrow beak.
Fully developed conidia of the neotype fit this
description well. Juvenile conidia ca. 50 x 10 wm with 4
transepta elongate but remain relatively narrow while a beak
as long as 90 wm is produced. The conidium body continues
to lengthen, expand, and lay down numerous transverse and a
few longitudinal septa; the beak lengthens, becomes septate,
and occasionally becomes 1-branched. No surface ornamenta-
=< 63 eee. Be :
ry
ee :
Sra omes ey = Sen f
see :
Z ee ‘
SN
|
- §
uo
ie
Ss
5
240
tion is evident on the conidia. The taxon is an excellent
representative of a large-spored, filament-beaked Alternaria,
to which genus the epithet is transferred.
Alternaria acalyphae (Nelen) E. Simmons, comb. nov.
Basionym: Macrosporium acalyphae Nelen, Bot. Mater.
Otd. Sporov. Rast. Bot. Inst. Akad. Nauk SSSR 15: 142.
1962. Neotype specified above.
A specimen of A. acalyphae was collected by H. C.
Greene in Wisconsin in 1948 (WIS, EGS 41-060). He reported
it as Alternaria sp. on Acalypha virginica L. in one of his
series of Notes on Wisconsin Parasitic Fungi (1949), from
which it was compiled in Farr et al. (1989). The conidia of
the Wisconsin specimen are for the most part juvenile or
young-mature filament-rostrate, rarely with a branched beak.
Otherwise the material is typical. No isolate of this fungus
is known to me.
88. 1962 Stemphylium botryosum Wallr. var. majus Batista
& Bezerra [as major], in Batista, Peres and Bezerra,
Inst. Micols. Univ.* Recite Publ: 343) 0p. 17), | Sobre
Croton campestris St. Hil... .lpojuca, Pern. Leg. Dr
Epaminondas de Barros Correia, 5/1/6l. Fipoye cis
IMUR,
The varietal epithet is illegitimate through lack of
a Latin description or diagnosis. The specimen that would
be considered type if the varietal name were legitimate (URM,
EGS 16-087) bears scarce, echinulate Stemphylium conidia up
to 48 x 22 wm (the authors give 27-43 x 12-19 wm) with 3-4
constricting transverse septa. Judged on the basis of the
few conidia and conidiophores seen and the authors’ illustra-
tion,.,.the. taxon. jis 7similar to.S.\,lycopersicin (Enjoja)
Yamamoto but certainly not to S. botryosum.
The variety has no nomenclatural standing and, due to
the scarcity of spores, is unidentifiable more closely than
to genus.
69" 1970 Stemphylium floridanum Hannon & Weber var.
euphorbiae Nag Raj & Govindu
Sydowia 23(1969): 115. On living leaf of Euphorbia
sp., Hebbal, Bangalore, 2.5.1966, T. R. Nag Raj.
241
Type (according to authors): MYSP + 823), with culture
from type also deposited in Herb. MYSP. I have not seen
either one of these.
The taxon, as described and illustrated by the authors,
has conidia "pale to olivaceous brown, oblong, cylindrical
or subangular, rounded or acutely pointed at the apex, often
with a beak, base truncate or rounded with a prominent basal
scar, 3-4 major horizontal septa at which the walls are
constricted and many secondary septa, walls verrucose, 41-69
x 14-19 uw in size, usually three times longer than broad."
The conidia described and illustrated are almost
certainly recognizable as S. lycopersici (Enjoji) Yamamoto
(syn. = S. floridanum Hannon & Weber) or a variety thereof.
904 1980 Alternaria crotonis Kamal, Singh & Kumar
Indian Phytopath. 32(1979): 228. On living leaves of
Croton bonplandianum Baill (Euphorbiaceae) ; University,
Gorakhpur; August 1976.
Type. (according to authors’); IMI. 206233, leg. R. P-
Sinenes Lo; University, Gorakhpur, U.P? , “India. I have=not
found this specimen in the herbarium at IMI; no other
specimen of the taxon, as named, is known to me.
The authors describe and illustrate a relatively small-
spored fungus, presumably an Alternaria, with solitary,
rarely catenate, obclavate, olive brown, smooth conidia 7.2-
45 x 3.6-10.8 wm (usually 21.6 x 7.2ym), with (in illustra-
tion) 3-4 transverse and rarely 1 longitudinal septa; the
tapering apex of the conidia, a beak, is simple, smooth,
hyaline, non-septate, up to ca. 20 zm in length.
The authors conclude that a novel species is in hand
because "it shows some morphological resemblance" to but
differs in one way or another from A. cheiranthi (Lib.)
Bolle, A. dianthi Stev. & Hall, A. tenuissima (Kunze ex
Pers.) Wilts., and A. ricini (Yosh.) Hansf. The comparisons
are spurious in that the conidia, as illustrated, have no
species resemblance whatever to the taxa listed. A. crotonis
is unknowable unless the type specimen becomes available.
oT. 1981 Ulocladium manihoticola Yen
Bulle Soc. Mycol srrance 9/152) snap itac an foi is
242
vivis Manihotis utilissimae in Sungei Way, Kuala
Lumpur, Malaysia, ad Jo-min Yen, No. 71172, 1 Oct. 1971
(Yen Herb. No. 10573).
Type (according to Yen): Yen Herb. No. 10573, leg. Jo-
min Yen, No. 71172, 1 Oct. 1971. Repeated attempts to borrow
this and other Yen specimens for examination have been
unsuccessful. Information from his institutional connection
at the time of his death (LAM) is that his herbarium is in
the hands of his family and unavailable.
The taxon, as illustrated, would be difficult to place
in Ulocladium but very easy to place in Pithomyces. Only
examination of the type specimen can yield an identification
of the taxon.
epiaiy 1983 Embellisia conoidea E. Simmons
Mycotaxon 17: 226. On old crustose, scarcely plastic
natural latex of cultivated Hevea sp.
Holotype: BPI 431684, on natural latex of Hevea sp.,
Java, Indonesia, Feb. 1969. EGS 29-179.
This taxon, with its brown, septate, conoid conidia 13-
15 x 5-7 ym produced individually at the apex of progressive-
ly geniculate conidiophores, probably would at one time have
been considered a rather odd, small-spored Alternaria. The
decision to segregate somewhat similar taxa with thick,
densely pigmented secondary septum layers abutting the paler
outer wall of the conidium into a separate genus Embellisia
E. Simmons (1971) gave a morphologically suitable systematic
placement for E. conoidea.
The species has no significant resemblance to any other
taxon treated here and is included only because the substrate
of its type is latex of a euphorbiaceous plant.
93. 1985 Alternaria euphorbiae (Barth.) Aragaki & Uchida,
in Yoshimura, Uchida and Aragaki
Plant Dis 70 C1986). 75.
Basionym: Macrosporium euphorbiae Bartholomew, Fungi
Columbiani no. 2633, iss. 1908, with type description on
printed label of exsiccatus packet. Lectotype: from BPI
0445288, as specified above in Alt.T&V 78. See Alt.T&V 78,
243
Fig. 71 for BPI lectotype conidia and Alt.T&V 97, Fig. 76 for
a more comprehensive range of conidia from the NY unit of F.
Col, 2633.
The observations and conclusions given in Alt.T&V 78
for M. euphorbiae Barth. need not be repeated here. Typical
A. euphorbiae has relatively large conidia which might be
thought of as belonging to an A. cucumerina or an A. zinniae
morphology group, i.e., with a long ellipsoid or ovoid spore
body and a long, narrowly filamentous beak.
Typical A. euphorbiae conidia reach 65-90 x 16-18 um
(spore body) plus a beak 120-150 x ca. 2 4m or less. The
conidium wall is smooth; both the outer wall and the trans-
verse septa are thin but well defined, with the transverse
septa showing a clear, medium golden brown color. Largest
conidia have 7-10 transverse septa and 1 or rarely 2 longitu-
dinal septa in 1-5 of the broadest transverse segments.
There is no evidence of apical secondary conidiophores, and
therefore of chain production, among conidia of the lecto-
type.
Alternaria euphorbiae has been seen in very few
specimens other than the Bartholomew F. Col. 2633 exsiccati.
I know of no isolate of this species.
Interestingly enough, a Kellerman and Swingle specimen
#1076, collected on Euphorbia marginata at Manhattan, Kansas,
Sept. 4, 1887, carries excellent large-spored Alternaria
material. A portion of this collection in BPI 0445524 (EGS
41-105) has abundant conidia of young-mature, filament-beaked
A. euphorbiae. Another portion of the same collection #1076
in KSC (EGS 06-140) has abundant conidia of strongly pseudo-
rostrate A. euphorbiicola Simmons & Engelhard (Alt.T&V 95).
Both portions of this collection also carry conidia of a very
distinctive Stemphylium of the S. solani Weber group. The
packets of both portions of the Kellerman and Swingle
collection are labeled "Macrosporium truncatum E. & K.," a
name that never was published and that is given here only as
an item of historical interest.
94. 1986 Alternaria euphorbiae (Barth.) E. Simmons & S.
Francis, in Simmons
Mycotaxon) 254195.
Basionym: Macrosporium euphorbiae Bartholomew, in
244
Fungi Columbiani no. 2633, iss. 1908, with type description
published on exsiccata label. This name was published in
Alternaria two months after Aragaki and Uchida (1985) had
published the same new combination derived from the same
basionym. The Simmons and Francis combination is an illegit-
imate later homonym.
However, each pair of combining authors chose a dif-
ferent unit of F. Col. 2633 as the source of their lectotype
fungus for the name M. euphorbiae Barth. Given the diversity
of Alternaria taxa present in the Aragaki and Uchida lecto-
type slide (see Alt.T&V 78), it is understandable but unfor-
tunate that each pair of authors chose a different large-
spored, filament-beaked Alternaria as _ lectotype. Both
species are present in units of F. Col. 2633; they are
readily distinguishable from each other, especially when
examined microscopically side by side. Working strictly
within ICBN rules on lectotypification, the Aragaki and
Uchida choice of lectotype must be maintained, as explained
in Alt.T&V 78. The Simmons and Francis lectotype choice
becomes irrelevant to the characterization of A. euphorbiae.
Ironically, both pairs of combining authors managed to
introduce elements of confusion into the systematics of
Alternaria species on Euphorbia species. Yoshima et al.
(1985) attributed an Alternaria disease of poinsettia to A.
euphorbiae, but the fungus they isolated, described, and
illustrated from blighted plants is a morphologically
different species whose conidia are not present in the
lectotype slide of A. euphorbiae. (See A. euphorbiicola,
Alt.T&V 95 below. )
Simmons and Francis (Simmons 1986) chose as lectotype
a species whose large, filament-beaked conidia differ in
width, pattern and definition of septa, and beak length from
lectotypical A. euphorbiae. Conidia of this as yet unnamed
species are present in F. Col. 2633 units at BPI 0445288
(Aragaki and Uchida slide #1; EGS 41-116), NY (EGS 03-047,
41-126), and PUR (EGS 41-128). The same species is known
from collections from other countries and from a Florida
isolate received for opinion in 1986. This material is used
as the basis of a new species proposal in Alt.T&V 98 below.
eye 1986 Alternaria euphorbiicola E. Simmons & Engelhard,
in Simmons
Mycotaxon 25: 196.
245
An avowed substitute (nomen novum) for Macrosporium eu-
phorbiae Reichert (1921) non M. euphorbiae Bartholomew
(1908). Holotype: B, auf Euph. prunifolia, Salamum prope
Mansurah., .6.12.i1911. SS. Schweinfurth .CEGS. 09-103). See
Alt. T&V 80 above.
Alternaria euphorbiicola is the correct name for a
species with erostrate to short taper-beaked conidia that
very frequently become strongly pseudorostrate, both in
nature and in culture. The conidia of this taxon do not
produce the narrowly filamentous beaks characteristic of A.
euphorbiae conidia.
Alternaria euphorbiicola is known from several species
of Euphorbia (detailed below), including E. pulcherrima
(poinsettia). A commercially important disease of poinsettia
was considered in detail by Yoshimura et al. (1985), where
the fungus was misidentified with Bartholomew's M. euphorbiae
(i.e., A. euphorbiae), and in passing by Simmons and Engel-
hard (Simmons 1986), where the fungus was given a new name
to replace the homonym M. euphorbiae Reichert.
Alternaria euphorbiicola is known not only from the
type specimen from Egypt (B, on E. prunifolia; EGS 09-103)
but also from Australia (BPI 0445274, on Euphorbia sp., USDA-
PPQ intercept Oct. 1962; EGS 41-104), Hong Kong (IMI 118473,
on E. pulcherrima, March 1966; EGS 26-110), Tanganyika (IMI
106673a, on E. grantii Oliver, February 1964; EGS 26-108),
USA-Florida (Engelhard 6386, 6392, 6468 isol. from E. pul-
cherrima; EGS 38-080/-081/-082), USA-Hawaii (Aragaki 1090,
1091, isol. from E. pulcherrima; EGS 41-028/-029) , USA-Kansas
(BPI 0446300, on E. marginata, Bartholomew, July 1899; EGS
41-094), USA-Kansas (KSC, on E. marginata, coll. Kellerman
and Swingle no. 1076, Sept. 1887; EGS 06-140), USA-Louisiana
(H. L. Walker, isol. from E. heterophylla L., EGS 38-131, and
from E. hyssopifolia L., EGS 38-191).
The conidium body length of both A. euphorbiicola and
A. euphorbiae may reach 80-90 um, but the width of the former
reaches 18-20 yum (in type) whereas that of the latter reaches
only ca. 15ym (in lectotype). Conidia of A. euphorbiicola,
essentially beakless, have an altogether more robust appear-
ance than do those of A. euphorbiae. In addition, sporula-
tion of A. euphorbiicola in nature is decidedly catenate by
means of sturdy apical secondary conidiophores, whereas A.
euphorbiae with its very long, sinuous, filamentous beaks
gives no evidence of secondary conidium production in the
246
natural specimens examined. Isolates of A. euphorbiicola
sporulate prolifically in culture, where colonies on PCA
produce conidia that commonly reach a size range of 88-112
x 29-32 ym before secondary conidiophores develop. Secondary
conidiophores, usually ca. 10-15 yum but often 28-48 um long,
have a diameter of ca. 4.5 ym until they expand into a
conidiogenous tip cell ca. 5.5ym diam. (And see Alt.T&V 15,
Fig. 28; Simmons 1986.)
96. 1986 Alternaria angustiovoidea E. Simmons
Mycotaxon 25: 198. On lesions of Euphorbia esula L.,
Manitoba, Canada.
Holotype: BPI 0445899, dried culture ex isol. K.
Mortensen 82-53, 27 July 1982, Manitoba, Canada (EGS 35-172).
This pathogen of a field pest, leafy spurge, is known
as isolates from Iowa and North Dakota, U.S.A., as well as
from Canada. It has little resemblance to any other taxon
on Euphorbia species described through 1986, in that its
heavily ornamented, usually narrowly ovoid to ellipsoid, long
pseudorostrate conidia are produced in moderately long,
usually unbranched chains of 8-10 units. Conidia 50-75 x 10-
15 wm in size often produce apical secondary conidiophores
in a 50-75(-100) x 4-5 ym range. (And see Alt.T&V 16;
Simmons 1986.)
Elam Bartholomew’s Fungi Columbiani 2633
and Some of Its Alternaria Contents
Stevenson (1971) reports that Bartholomew proposed to
issue 70 copies of each Century of Ellis and Everhart’s Fungi
Columbiani when he took over as editor and publisher of this
exsiccata with Century XVI in 1901. (The full name had been
reduced simply to Fungi Columbiani.--E. Bartholomew on
specimen packets by the time no. 2633 of Century XXVII was
issued in 1908.)
I have had the opportunity to examine the contents of
six packets of F. Col. 2633, each with its original printed
label carrying the type description and collection data of
Macrosporium euphorbiae Bartholomew. The six packets are in
BPI 0445288, 0445289, 0445290 (EGS 41-116, 41-117; 41-118);
NY (examined twice, EGS 03-047 and 41-126); PUR (EGS 41-128) ;
and WIS (EGS 01-196).
247
The 3-4 Euphorbia leaves in each packet are in good to
excellent condition. Each leaf has several small, brown,
round lesions, as described by Bartholomew, some of them
obviously covered densely with fungus growth but others not
conspicuously so. A few leaves also have 1-2 small fungus-
bearing lesions that are somewhat irregular rather than quite
circular in outline. A cursory macroscopic examination gives
an impression that the packet contents are "duplicates," in
the sense of appearing to be portions of one large collection
from a single host colony, or at least of several specimens
of one host species, all exhibiting the same kinds of
leafspot lesions.
Examination of even an extremely limited sampling of
fungus material from leaf lesions in each packet (at 195X and
780X) invites a state of alternarial shock or at least
surfeit from the presence of at least four quite distinct
Alternaria taxa. An inevitable cautionary note comes to
mind, to the effect that neither the specialist nor the tyro
mycotaxonomist should expect that leafspots collected in
summer and autumn of Midwest U.S.A. (or anywhere, for that
matter) will bear pure stands of single fungus species or
that different lesions on a leaf will necessarily bear the
same fungus taxon.
This observation, the diversity of conidium morphology
observable on the Macrosporium euphorbiae lectotype slide
(see Alt.T&V 78 above), and the published differences of
opinion on the identity of the taxon have suggested a review
of slide preparations from units of F. Col. 2633 that have
been readily available to me.
WIS (EGS 01-196) My first encounter with a unit of F. Col.
2633 was disappointing, in that an Alternaria taxon present
could not be identified with the characterization presented
in the accompanying type description. The obvious Alternaria
present is a taxon that can be found on a high percentage of
leafspots of herbaceous and woody plants in temperate
climates (cfr. Alt.T&V 46, Fig. 51; Simmons 1990).
NY (EGS 03-047) A second encounter with F. Col. 2633 yielded
conidia of a fungus that determined my concept of the taxon
until 1986, when Simmons and Francis presented it as their
choice of lectotype of M. euphorbiae Barth. and proposed
Alternaria euphorbiae (Barth.) E. Simmons & S. Francis, comb.
nov. This fungus is discussed in Alt.T&V 14 with Fig. 27
(Simmons 1986). It is a large-spored, filament-beaked
248
Alternaria that matches part but not all of the type charac-
terization. With the experience of having examined several
portions of F.. Col. 2633, , it is safe. to say that’ this taxon
is not A. euphorbiae, although it is present in several units
of this exsiccatus. Its choice as lectotype of M. euphorbiae
Barth. was preceded by the Aragaki and Uchida lectotypifica-
tion (Yoshimura et al., 1985), as explained in Alt.T&V 78.
As a distinct, as yet nameless Alternaria, it is described
below as a new species on the basis of modern material (Alt.
T&V 98).
BPI 0445288 (EGS 41-116) An identification problem (early
1993) with a quite different, at first sight novel Alternaria
from Euphorbia esula in Manitoba and Saskatchewan, Canada,
plus the need to clarify the identity of A. euphorbiae on E.
marginata and A. euphorbiicola on E. pulcherrima, lured me
into reviewing slides from some 130 specimens of Alternaria,
Macrosporium, Stemphylium, etc., from substrates of Euphorbi-
aceae held mostly at BPI and IMI.
Among BPI holdings are three packets of F. Col. 2633.
Each of the three is stamped TYPE. One, BPI 0445288, is
accompanied by a microscope slide preparation marked #1,
representing the evidence of lectotype choice made by Aragaki
and Uchida. As mentioned above (Alt.T&V 78) conidia of three
different and definable morphologic taxa of Alternaria (and
a probable Nimbya) are present in this slide preparation.
I made only one additional slide preparation from this
BPI unit of F. Col. 2633, from one lesion on which conidia
could be seen at 50X magnification. A fourth Alternaria
taxon, remarkably different from the other three, is present.
Each of the four taxa is discussed in turn in Alt.T&V 97-100
below.
ihe Alternaria euphorbiae (Barth.) Aragaki & Uchida
Fig. 76
See Alt.T&V 78 and 93 above for characterization and
illustration of the lectotype in BPI 0445288. Conidia of
this taxon present in a second unit of F. Col. 2633, BPI
0445290, are either old with lateral germination tubes or
juvenile, presumably representing a secondary flush of
sporulation on the lesion. No conidia of A. euphorbiae were
found in a preparation from a single likely-looking spot
(among ca. 15) of a third unit of F. Col. 2633, BPI 445289.
249
Fig. 76. Alternaria euphorbiae: conidia and conidiophores
from NY unit of Fungi Columbiani 2633 (Macrosporium euphor-
bDiaeesarch.)..) *Bar == 50721
250
Young-mature to fully developed conidia of A. euphor-
biae are abundant on lesions of two leaves (of four) in the
NY unit of F. Col. 2633 (EGS 41-126). Excellent conidia of
the species in all stages of development also are abundant
on lesions of all four leaves of the PUR unit of the same
exsiccatus (EGS 41-128). The WIS unit provided no A.
euphorbiae conidia in the single slide preparation obtained
(EGS 01-196).
Figure 76 allows presentation of a wider range of A.
euphorbiae conidium development than is visible in the
lectotype slide. Euphorbia marginata and A. euphorbiae
probably still flourish in the neighborhood of Stockton,
Kansas (the type locality); modern air-dried specimens
suitable for isolation procedures would be received with
gratitude.
98. Alternaria obtecta Simmons Fig.
Alternaria sporulation elements present on some lesions
of the F. Col. 2633 unit at NY (EGS 03-047) were character-
ized and illustrated by Simmons and Francis in their discus-
sion of M. euphorbiae Barth. (Simmons 1986). The fungus is
one of two Alternaria taxa with large, filament-beaked
conidia that occur on this specimen. Typical A. euphorbiae,
as discussed above in Alt.T&V 74 and 98, is one of the two
taxa. The other one, as treated by Simmons and Francis, is
not A. euphorbiae, although, on the basis of the NY specimen,
they considered it to be so at the time. It is, instead, a
distinguishable species known from other collections and from
an isolate which came to my attention in Nov. 1986. It is
described here as novel.
Alternaria obtecta E. Simmons, sp. nov.
Etym.: L. obtecta, concealed, covered over, with
reference to the obscurant quality of the fine spore
ornamentation
Ex cultura in agaro PCA descripta. Conidiophora
simplicia vel geniculata, dilute fulva, 30-80 x 4-6 ym.
Conidia' solitaria, raro lin \catenis bisporis. Conidia
longiovoidea vel ellipsoidea, dilute fulva, usque ad 65-70
x 16-19 wm, rostro recto vel flexuoso ad 110-160 x 1.5 zm,
transverse 10-12 et longitudine 2-5 septata, episporio sub-
tiliter et obtecte punctulato. Habitatio typi in maculo
Euphorbia supina Raf., Ruston, Louisiana, U.S.A. Holotypus:
2a
“oe hey
te at) 59
ee ae
te
Fig. 77. Alternaria obtecta: conidia and conidiophores from
holotype (BPI, EGS 38-189); all conidia punctulate to ex-
tremely minutely ornamented. Bar = 50 wm.
202
pars ex cultura EGS 38-189 desiccata et in BPI conservanda.
Narrowly ellipsoid or ovoid juvenile conidia ca. 40-50
x 10-12 wm have 6-7 transverse septa without evidence of
longisepta, a condition that holds as the conidium body
broadens, lengthens, and initiates an apical filamentous
beak. Even the youngest conidium walls are completely and
densely punctulate. The ornamentation persists throughout
the development of the spore, as distinct punctulation (at
780X) or as an almost imperceptibly fine covering. The layer
of ornamentation obscures the few longitudinal septa to such
an extent that the evenly spaced transepta appear (at 195X)
somewhat ladder-like. Three or four of the 10-12 transepta
in largest conidia are darker and somewhat more constricting
than the others; one longiseptum can be found in each of 2-5
of the transverse segments. Ornamentation is less pronounced
and obscurant on conidia produced on V-8 agar.
Fully developed conidia are long ovoid or ellipsoid,
reaching a size range of 65-70 x 16-19 wm and tapering into
a straight to),filexuous’ beak (upto! 110-160 x ca.2jl.oeym:
Conidium color is a pale golden tan. Evidence of chain
formation is present, though rare, with the tip of a filamen-
tous beak enlarging to about 10 x 4 ym and becoming conidi-
ogenous.
The holotype was prepared from an isolate received
November 1986 from L. Walker, Ruston, Louisiana, as "231
Alternaria sp.," from prostrate spurge (Euphorbia supina).
(The isolate is the property of Mycogen Corporation, San
Diego, California.) Conidia of A. obtecta are found also on
some lesions of E. marginata leaves in F. Col. 2633 (BPI
0445288, NY, PUR) in association with A. euphorbiae, A. con-
stricta, and various small-spored Alternaria taxa.
99, Alternaria sp.
The small-spored, chain-forming Alternaria that occurs
on the lectotype slide of M. euphorbiae Barth. along with
conidia of A. euphorbiae and A. obtecta is illustrated above
invAlte.T&V 07838 Fie.) 72, fori contrasts purposes...) Conidiagor
this taxon are readily distinguishable from those of the
other two species. However, too few conidia are present to
permit its identification with other small-spored taxa. Most
of the conidia are narrowly ellipsoid with either a gradually
attenuate true beak or a short, sturdy secondary conidio-
253
phore. The largest conidium bodies are dilute ochraceus in
color, smooth or finely but distinctly punctulate roughened,
ca. 40-45 x 10-13 wm plus a beak ca. 18-20 x 3 ym or an
apically perforate secondary conidiophore ca. 15-18 x 4 yum.
100. Alternaria constricta Simmons Fig. 78
A fourth distinctive Alternaria species that occurs on
some portions of Bartholomew's F. Col. 2633 was noticed first
in the single slide preparation made by EGS from BPI 0445288
(EGS, 41-129)..." This’ is the same unit of @F.\Col, 2633 ‘that
furnished the lectotype of M. euphorbiae Barth. in slide #1
of Aragaki and Uchida, although conidia of this fourth taxon
have not been seen on the lectotype slide itself.
Fortunately, good material of this fourth taxon also
is found in the NY unit of F. Col. 2633 (EGS 41-126), which
can be used as its holotype.
Alternaria constricta E. Simmons, sp. nov.
Etym.: L., constricta, a diagnostic conidium character
Ex specimine siccato descripta. Conidia solitaria vel
interdum in catenis. Conidia breviellipsoidea vel ovoidea,
ad aliqua transepta primaria distincte et fortiter constric-
ta, erostrata aut Jlongifilamentose rostrata, interdum
pseudorostrata et conidifera; dilute sordide fulva, plerumque
laevia, 40-65 x 13-18 wm, transverse 4-7 et longitudine 1-3
septata, rostro filamentoso ad 100(raro -150) x 2-3 zn,
pseudorostro ca. 40 x 54m. Habitatio typi in maculo folii
Euphorbia marginata Pursh, Stockton, Kansas, U.S.A. Holo-
typus: NY, praeparatio permanens lamina vitrea EGS 41-126
ex specimine Fungi Columbiani exs. no. 2633 (non raro alliis
speciebus commixta), leg. E. Bartholomew, July 20, 1908.
Beakless conidia and the spore body of rostrate conidia
are short ellipsoid or ovoid, distinctly and sometimes
strongly constricted at 2 or more major transepta. Conidia
apparently may remain beakless, but most produce an apical
filamentous beak. Occasionally a conidium extends into a
sturdy well-defined secondary conidiophore, or the tip of a
filamentous beak enlarges and becomes converted into a
functional secondary conidiophore.
Conidium bodies and beakless conidia are dilute dull
golden tan in color, apparently quite smooth but occasionally
254
Fig. 78.
Alternaria constricta:
EGS 41-126). Bar = 50 pm.
conidia from holotype (NY,
25
with small scattered verrucae, 40-65 x 13-18 wm, with 4-7
darker transverse septa and 1-2 thin, longitudinal or oblique
septa in 1-3 of the transverse segments. Filamentous beaks,
which commonly arise quite abruptly without tapering from the
conidium apex, are straight to slightly curved, and up to 100
(rarely to 150) x 32-3 pm.
The holotype consists of conidia present in a permanent
slide preparation EGS 41-126 derived from a single lesion
present on a leaf of Euphorbia marginata in the specimen of
Fungi Columbiani no. 2633 in NY. The individual leaf and
lesion from which the holotype was prepared are marked for
reference.
Additional Alternaria Taxa on Euphorbiaceae
101. Alternaria agripestis Simmons & Mortensen ie oven Ae)
Euphorbia esula (leafy spurge), like E. marginata, is
host to more than one strikingly distinctive Alternaria
taxon. K. Mortensen was the original isolator of the type
of A. angustiovoidea E. Simmons (1986) from E. esula in Mani-
toba. His recent sharing of isolates, field information, and
supporting evidence of pathogenicity of this additional Al-
ternaria from E. esula motivated this present review.
Alternaria agripestis E. Simmons & K. Mortensen, sp. nov.
Etym.: L. ager + pestis (gen.) = of a field pest
Ex cultura in agaro PCA descripta. Conidiophora
simplicia vel geniculata, sordide fulva, ca. 80-225 x 6-8 ym.
Conidia solitaria, raro in catenis bisporis, longe angusti-
ovoidea vel anguste ellipsoidea, sordide fulva, usque ad 80-
100 x 16-18 “zm, rostro inramoso 200(-250) “zm longo x 3-4 “zm
a 1.5 wm decrescenti vel rostris 1-3-ramosis 100-200 zm
longis, transverse 9-11 et longitudine 1-5 septata, laevia,
punctulata, vel fortiter verruculosa. MHabitatio typi in
maculo caulis Euphorbia esula L., Maxim, Saskatchewan,
Canada. Holotypus: pars ex cultura EGS 41-034 desiccata et
in BPI conservanda.
Well-developed conidia of A. agripestis, like those of
A. angustiovoidea, are strikingly long-narrow ovoid with
numerous transverse and very few longitudinal septa.
Conidium walls of A. agripestis remain smooth to sparsely
256
punctulate or become strongly verrucose under some culture
conditions (see below), whereas those of A. angustiovoidea
typically become densely verrucose. Here the resemblance of
the two ends, as A. agripestis conidia are solitary with a
long narrow beak which often is branched whereas A. angusti-
ovoidea conidia are produced in long, usually unbranched
chains of 8-10 pseudorostrate units.
Juvenile conidia of A. agripestis are narrowly ovate,
gradually tapered to a pointed beak, and as large as 50-60
x 8-9 wm when 4-5 transverse septa become visible. Conidia
in this stage of growth are subhyaline but already visibly
punctulate. Development of long narrow beaks is rapid as the
conidium body grows in length and somewhat in width; single
long beaks are the common condition on well-developed
conidia, once-branched beaks are abundant, and twice-branched
beaks are present though rare.
Largest conidia have bodies in the 80-100 x 16-18 yum
size range, are dull golden tan in color, 9-11 transversely
septate with one longitudinal or oblique septum in each of
1-5 transverse segments. Solitary unbranched beaks are
straight to smoothly curved, subhyaline, up to 200(-250) um
long, tapering from a base 3-4ym wide to a fine 1.5 wm apex.
Chains of two conidia occur, though very rarely.
The above type description is based on 7-day PCA cul-
tures produced under routine conditions outlined in the
introductory comments. Under these conditions sporulation
is tardy and sparse. However, abundant sporulation on PCA
and enormous numbers of conidia on V-8 can be induced within
24 hrs. by means of an old plate-culture trick, viz., scari-
fication of the colony surface followed by a few hours of
fluorescent-light exposure and overnight hours of darkness.
Conidia of A. agripestis produced using this method of
culture manipulation are quite similar in size (ca. 72-88 x
14-16 wm) and are densely verrucose in ornamentation, which
is concentrated on the lower half to two-thirds of the spore
body. Beak branching becomes the rule, with once-branched
beaks dominant, twice-branched (3 beaks) common, and thrice-
branched (4 beaks) present though rare.
The holotype is from an isolate received 26 January
1993 from K. Mortensen, Regina, Saskatchewan, as a 1-conidium
isolate 92-81A from infected stem tissue of Euphorbia esula
collected by P. Harris, Maxim, Sask., July 9, 1992.
pial
Fig. 79. Alternaria agripestis: conidia from holotype, 7d.
on PCA (BPI, EGS 41-034). Bars = 50 wm.
258
102. Alternaria subulata Simmons Fig. 80
M. B. Ellis, hyphomycetologist at IMI for many years
until his (semi)retirement, frequently shared with me speci-
mens of Alternaria taxa whose identities he did not intend
to pursue for one reason or another. He assigned each
problem taxon a "taxonomic species" number when depositing
examples in IMI. Some of these items are scanty, sometimes
only a scrap of host and a slide, and perhaps will never
prove usable for classification purposes. Others are quite
adequate for comparison work.
It seems pertinent to comment here that sometimes it
is difficult to judge whether or not a conidium sample from
a field specimen contains fully developed spores along with
obvious juveniles. Many small-spored Alternaria taxa can
produce fairly extensive chains of conidia overnight under
dampchamber conditions, even though the original field
specimen held only a flush of solitary, noncatenate spores.
Defining such a taxon on the basis of the field specimen is
misleading and troublesome to classification work, although
it has been done many times in the literature of the genus.
Two specimens of "Alternaria Tax. sp. XI" (MBE in IMI)
present such a dilemma. I have decided to present the taxon
here as novel because its specimens contain developmental
stages unusual enough for subsequent recognition and because
it occurs on a single host species, Euphorbia hirta L., in
collections made 11 years apart in two widely separated geo-
graphic regions, New Guinea and Sudan.
Alternaria subulata E. Simmons, sp. nov.
Etym.: L. subulata, awl-shaped; of the conidia, though
not so sharp-pointed as a hand-awl (= subsubulate?)
Ex specimine siccato descripta. Conidiophora simpli-
cia, geniculata, dilute ochracea, pluriconidifera, ca. 50-75
x 4-5 yum. Conidia solitaria, anguste ellipsoidea, dilute
ochracea, laevia, erostrata 35-50 x 10-13 um, transverse 4-5
et longitudine 0, raro 1 septata, vel rostro 50-65 x 4 ym.
Habitatio typi in maculo Euphorbia hirta, S. Staines, Gusap,
New Guinea. Holotypus: IMI 117496b (EGS 22-050), leg. D.
Shaw et W. Burnett, 20 January 1966.
Juvenile conidia are ellipsoid and beakless, ca. 25 x
7 um by the time 3-4 transverse septa have developed. Apical
beak growth is initiated at this stage; the beak lengthens
227
Fig. 80. Alternaria subulata: conidia and conidiophores
from holotype (IMI 117496b). Bar = 50 wm.
260
as the conidium body enlarges. Largest conidia are 35-50 x
10-13 wm, dilute yellowish tan, smooth, with 4-5 transverse
septa and rarely 1 longitudinal septum in one transverse
segment, and with a narrow but not truly filiform beak up to
50-65 x ca. 4ysm. There is no evidence of secondary sporula-
tion in either field specimen known for this taxon.
Holotype IMI 117496b is a portion of a_ specimen
collected by D. Shaw and W. Burnett (Terr. Papua and New
Guinea Dept. Agr. 4887) on leaf spots of Euphorbia hirta, S.
Staines, Gusap, New Guinea, 20 January 1966. A second
representative of the species in IMI 61966 (EGS 22-051) was
collected by B. Beshir (Sudan Mycol. Herb. 2793) on the same
host species, Zalingei, Sudan, 22 October 1955.
103. Alternaria acalyphicola Simmons ELgveou
A second Alternaria on an Acalypha species was sent to
me in 1985 as a culture from IMI. It is distinctly different
not only from A. acalyphae (Nelen) Simmons (Alt.T&V 87 above)
but also from any other Alternaria taxon described from a
euphorbiaceous substrate.
Alternaria acalyphicola E. Simmons, sp. nov.
Ex cultura in agaro PCA descripta. Conidiophora
simplicia vel ramosa, geniculata, dilute castanea, 65-100 x
5-7 ym. Conidia solitaria, raro in catenis bisporis,
juventute anguste ovoidea, copiose pustulate vel tuberculate
exasperata, maturitate punctulata, longe ovoidea vel ellipso-
idea, dilute castanea, usque ad 75-80 x 18-20(-22) um, rostro
inramoso filamentoso 100-175 x ca. 2 ym, transverse 8-10 et
longitudine ca. 5-7 septata. Habitatio typi in Acalypha
indica L., Seychelles. Holotypus: pars ex cultura EGS 38-
100 desiccata et in BPI conservanda.
Juvenile conidia usually are totally pustulate- or
tuberculate-roughened, the ornamentation nearly obscuring a
view of initial transverse septa. They commonly are ca. 40-
45 x 10-14 zm with 5-6 transepta when a narrow apical beak
begins to develop. The pustules become less crowded and
decreasingly prominent as the spore body enlarges, as though
they were being moved apart and pulled flat when the wall
surface area increases. Some fully developed conidia may
retain a scattering of pustules, but usually the mature
surface is only punctulate, not prominently ornamented.
261
Colt Oly 2 ?
A
ke GAO)
Fig. 81. Alternaria acalyphicola: conidia and conidiophores
from holotype (BPI, EGS 38-100); all juvenile conidia densely
ornamented, largest conidia almost smooth. Bar = 50 ym.
262
The spore bodies of fully developed conidia commonly
reach a size range of 70-85 x 18-20(-22) um plus a filamen-
tous, unbranched beak 100-175 x ca. 2 wm. Largest conidia
have 8-10 transverse septa and 1-2 longitudinal or variously
oblique secondary septa in 5-7 of the transverse segments;
they are a dilute chestnut brown in color with darker septa.
Evidence of secondary sporulation is rare, usually in the
form of a 1-cell, perforate swelling at the tip of a primary
beak.
The holotype was prepared following reisolation from
a culture received 10 May 1985 as IMI 266969 Alternaria sp.,
ex Acalypha indica, Seychelles.
104. Alternaria heveae Simmons Fig. 82
Two collections of Hevea brasiliensis leaves, gathered
a year apart in Mexico, bear sparse growth of a single Alter-
naria species. Conidia, though present on leaf lesions, are
difficult to locate, possibly because their wall color is so
dilute. The fungus is distinguishable from all other species
on Euphorbiaceae treated here. It is presented as a new
species, with hope that more abundant modern material
suitable for isolation work will be found.
Alternaria heveae E. Simmons, sp. nov.
Ex specimine siccato descripta. Conidiophora simpli-
cia, geniculata, 15-35 x 4-5 ym, subhyalina. Conidia soli-
taria vel brevicatenata, anguste ovoidea vel ovoidea, dilute
ochracea, laevia, transverse 4-5 et longitudine 1-2 septata,
60-95(-110) x 12-20 “zm (rostro incluso). Habitatio typi in
maculo Hevea brasiliensis, El Palmar, Mexico. Holotypus:
BPI 0445779 (EGS 41-087), leg. W. J. Martin, June 8, 1946.
Juvenile conidia are narrowly ovoid; the spore body
retains this shape as it enlarges and produces a gradually
narrowing, sturdy (not filamentous) apical beak. Conidia
have 6-7 transverse septa without producing longitudinal
septa, or 1 longitudinal or oblique septum may develop in 1-2
of the broadest transverse segments. The tip of some beaks
develops into a functional but morphologically scarcely dif-
ferentiated (slightly swollen, perforate) secondary conidio-
phore. Juvenile conidia are about 30-50 x 8-9 wm with 4-5
transverse septa before beak growth is evident. Largest co-
nidia are 60-95(-110) x 12-20um, dilute yellow-tan, smooth.
263
™
—1~
Fig. 82. Alternaria heveae: conidia and conidiophores from
holotype (BPI 0445779). Bar = 50 yum.
264
Both the holotype BPI 0445779 and a second specimen of
this taxon on Hevea brasiliensis were collected by W. J.
Martin at El Palmar (Veracruz), Mexico, the holotype on June
8, 1946, and the second BPI 0445781 (EGS 41-089) in May 1947.
105. Alternaria cicina Simmons Fig.) 33
Lesions in a collection of Ricinus leaves from Pakistan
have growth of an Alternaria species that brings to mind some
of the small, essentially beakless conidia of taxa discussed
in the first paper of this Alt.T&V series, e.g., A. mou-
chaccae E. Simmons (1981). Similar, probably conspecific
conidia occur also on a Ricinus leaf collection from Northern
Rhodesia (Zambia). In lieu of cultures, which are of special
value when characterizing relatively small-spored Alternaria
taxa, this fungus is described as novel on the basis of field
material produced under tropical but otherwise undefined
conditions.
Alternaria cicina E. Simmons, sp. nov.
Etym.: L. cicinus, adj. rel. to the castor-oil plant,
fr. Gr. kiki, the castor-oil plant (fide Brown, 1956,
who with Stearn, 1992, are our ever-present help in
time of Latin need)
Ex specimine siccato descripta. Conidiophora simplicia
vel ramosa et geniculata, dilute ochracea. Conidia solitaria
vel brevicatenata, plerumque anguste vel late ovoidea, vel
late ellipsoidea, erostrata, interdum obclavata, apice
conidifero, usque ad 30-45 x 12-17 4m, parce ochraceo, laevia
vel minute punctulata, transverse 5-8 et longitudine plurime
septata. Habitatio typi in maculo folii Ricinus communis,
Karachi, Pakistan. Holotypus: IMI 75883a (EGS 26-192), leg.
Azmatullah Khan (468), January 1958.
Juvenile and mature conidia alike are mostly narrow-
or broad-ovoid and beakless. Conidia may remain ovoid or
broad-ellipsoid with a bluntly rounded apex, or individual
conidia may elongate, taper somewhat distally, and produce
a small, thin-walled 1-cell secondary conidiophore. Well-
developed conidia are ca. 30-45 x 12-17 ym, with 5-8 complete
or partial transverse septa and 1-2 longitudinal or variously
oblique septa in most of the transverse segments. Conidium
color is a medium tawny brown with darker septa; walls are
smooth to minutely punctulate.
265
Fig. 83. Alternaria cicina: conidia and conidiophores from
holotype (IMI 75883a). Bar = 50 wm.
266
Material similar to the holotype was collected on the
same host species by A. Angus (434) at Mt. Mukulu Research
Station, Northern Rhodesia, 7 February 1958 (IMI 75692, EGS
25-115).
A note on other herbarium specimens of Euphorbiaceae
examined but not discussed above. This review has covered
examination of approximately 130 isolates and field specimens
purportedly of Alternaria, Stemphylium and somewhat similar
genera. Most of the novel or otherwise remarkable taxa are
included above. Many specimens contained Alternaria taxa of
small-spored, chain-forming kinds that for the most part
cannot be discussed profitably at present without using
cultured isolates.
ACKNOWLEDGEMENTS
Dr. Mary E. Palm yet again has shown her support of
this continuing series of Alternaria papers by providing
professional encouragement as well as essential, critical,
and much appreciated review of the prepublication manuscript.
The several institutional herbaria and culture collections
mentioned in the text have been generous with their loans and
gifts of material for study. Dr. M. Aragaki, University of
Hawaii, Honolulu, has shared some of his isolates from
Euphorbia pulcherrima. Dr. K. Mortensen, Agriculture Canada,
Regina, SK, Canada, with his continuing interest in pathogens
of weedy spurges, has supplied the original isolates not only
of A. angustiovoidea but also of the A. agripestis described
herein as novel.
Key to Alternaria Taxa Known on Euphorbiaceae
Any key to Alternaria taxa now known from euphorbiace-
ous substrates will be inherently distorted, in that some of
the species have been studied from natural collections,
others from cultures, but from both in only about half the
cases presented here. A good example of the hazard exists
in A. ricini, whose conidium shape and ornamentation in
culture are confusingly different from those observed in
natural collections from which isolates were derived. In the
following key a number preceding each epithet refers to an
Alt.T&V treatment of each species.
267
Key: Alternaria Species on Euphorbiaceae
Beak narrowly filamentous. .. . Sear B/b
Beak sturdy-tapered, pseudorostrate. or r Jacking ne Aa
B. Beak unbranched. .... ae a eee ene C/T
b. Beak branched (sometimes rare) Sis ane.) se ALGAE
Conidia smooth (in nature)... ees
Conidia ornamented (in nature endion palctreyy 3) eae hy 8
Dew conidia catenatve Cinsnature) “).tr., .6 28 ja eelB/e
OemCOnld a sOlicarya(ieneature init. Sine. ee ewe 2 a E/T
Transepta conspicuously constricting . . 100 constricta
Transepta only moderately constricting ... 83 ricini
F. Conidia ellipsoid to long ovoid. . 93,97 euphorbiae
£7 ~Conidiasnarrow ellipsoid) 2. 7. Ry 87 acalyphae
Beak branching rare (in nature). .... 87 acalyphae
Beak branching dominant, multiple. . . .101 agripestis
H. Conidia often catenate in culture. ... 83 ricini
Nneweconidia rarely \catenate in culture)... 2.0. /1/1
Ornamentation punctulate, becoming
Pine ObSCuLant 1) age, \.7es Sea age 98 obtecta
Ornamentation Pineda iiaee foerccul ates
becoming inconspicuous in age . . 103 acalyphicola
Jer vConi dia (solmrary tiie. ie Treas Be LOZ) subulata
He eCOntG ta (Catenace aan it. Wes. ae aes, Been ee RK
Conidia strongly ornamented. ... . 16 angustiovoidea
Conidia smooth in nature and in culture (if known). L/1
L. Conidia erostrate or weakly
pseudorostrate. ... ie Mois hOOeCicina
1. Conidia erostrate or Pourdidys pseudorostrate . M/m
Erostrate conidium apex short,
Hharrow-taperedst wha ens «est sth aa. 15 euphorbiicola
Erostrate conidium apex long,
SstuLdy=taperLed., . vole. 9. eteaw eee. . 5104 bheveae
268
LITERATURE CITED
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269
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270
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MY COTAXON
Volume L, pp. 271-278 January-March 1994
AGARICUS PSEUDOARGENTINUS N. SP. FROM ARGENTINA.
Edgardo Albert6é & Jorge E. Wright.
Departamento de Ciencias Bioldégicas,
Universidad de Bs.As., Ciudad Universitaria
(1428) Buenos Aires, Argentina.
Abstract: A. pseudoargentinus, a new
edible species found in the environs of
Buenos Aires, is described and
illustrated. It probably belongs to
section Sanguinolenti.
INTRODUCTION
During the Autumn months it is common to
find in the environs of Buenos Aires a great
diversity of species of Agaricus. They have been
recorded by Spegazzini (1899, 1926), Martinez,
(1949, 1957), Raithelhuber (1977, 1988) and
Heinemann (1986, 1987, 1990).
Recently, specimens of a species that does not
fit any described, were found and is here
proposed as a new taxon.
MATERIALS AND METHODS
Specimens were studied both macro- and
microscopically immediately after collection,
annotating their colours according to the charts
272
of Munsell [M] (1954) and Maerz and Paul [MP]
(1930) and performing upon them the pertinent
chemical reactions. The slenderness index (IG) as
used by Heinemann (1983) is calculated as IG:
L2/D.d, where L, D and d are stem length, pileus
diameter and stem diameter, respectively.
Cultures were also made in the fresh
condition by inoculating slant tubes containig
potato-glucose agar (Ferri, 1985) with portions
of the flesh. The abbreviation BAFC stands for
the mycological herbarium of the above
Department.
Agaricus pseudoargentinus Alberto et Wright n.
sp.
Pileo usque ad 60-110 mm diam., primo
gZloboso-haemispherico, dein convexo, fibrilloso
vel fibrilloso-squamoso, squamulis obscuris
densissimis ad disco centralis, fibrillis
centralis obscuriorum, curtis, tenuis, delicatis,
ad margo secedentibus; sub fibrillis niveo vel
Pallide brunneis, margo ab initio convoluto,
integro, dein plano, appendiculato, breviter
excedente supra lamellis. Superficies pileo ab
hyphis hyalinis vel castaneis, cylindricis, 7-15
um diam. Stipite 40-130 x 10-20 mm, cylindraceus,
niveus, fistulosus, fibrosus, in substrato parte
basali rhizomorphis efformantibus. Annulo
centrali vel superiori, simplex, membranaceo,
descendente, pendulo niveo, persistente in
maturitate. Lamellis liberis, confertis,
ventricosis, acie integro, primo cinereis, dein
pallide brunneis, in maturitate obscure brunneis.
Pulvis sporarum obscure brunneis. Contexto ad
centrum 10-17 mm, carni nivea. immutabilis, odori
fungoso dulci, sapore grato. Edulis.
Sporis: .4.7-5.7(6.5). xX: (322)3.70-4.2 un
curte ellipsoideis, brunneis, sine poris apicalis
cum apiculo discernibile. Basidiis tetrasporis,
claviformibus, 20-22 x 6-7 um; sterigmatis curtis
usque 3 um long. Cheilocystidiis brunneis, late
273
piriformibus vel globosis subglobosis, 16-25 x
09-14 um .
Reach Vochaertrers Wie ieen hah Sw ia =) vecido
sulphurico: Jlaeve; acido nitrico: laeve. Hab:
solitario, sed abundantibus inter herbis, post
pluvias.
Holotypus: Argentina, Bonariae, Burzaco,
leg. EF. Alberté. 2-V-1991, in Herb. BAFC ng
32.765 conservatus est.
Fig.1: A.pseudoargentinus n. sp. a- Habit. b-
Detail of the pileus surface showing the brown
fibrils.
274
Pileus 60-110 mm diam., when young globose-
hemispheric, later convex, with abundant brown to
Gark brown fibrils or fibrosquamules (10YR 6/6
[M]) with a darker central disk due to a higher
density of fibrils, which are then shorter,
thinner and more delicate, later separating
towards the margin, and becoming longer and
thicker (Fig. 1b),between white or tea-and-milk
colour (11 C/5 [(MP]); margin at first convolute
and smooth, later plane, appendiculate, entire,
slightly exceeding the lamellae. Stipe 40-130 x
10-20 mm, cylindric, white, sometimes with a
brownish hue at the base, hollow, fibrous,
forming rhizomorphs at the base inside the
substrate. Annulus central or superior, simple,
membranous, and descending, pendulous, skirt-
like, white, persistent at maturity (Fig. la).
Lamellae free, crowded, ventricose, with smooth
edges, at first white to greyish, later tea-and-
milk coloured, passing through sepia and finally
dark brown. Spore print dark brown (56 J/2),
difficult to obtain. Context in the centre of
fruit-body 10-17 mm thick, white, unchangeable;
odour mild, slightly fungous; taste pleasant,
Similar to A.bisporus. Edible.
Spores: 4.7-5.7(6.5) = (3.2)3.70-4.2 ium
(Q:1.25-1.45) n= 30 shortly elliptic; smootn:
brown, without an apical pore and with a visible
apiculus. Basidia 4-spored, clavate, 20-22 x 6-7
uM 5 sterigmata short, (byan haere, 2 jum. long.
Cheilocystidia: hyaline to brown, widely pyriform
or subglobose, 16-25 x 09-14 um (Fig.2).
Pileus covering of hyaline and_ brown,
cylindric hyphae, 7-15 um diam.
Aspect slender, IG: 2.7-8.5 (Heinemann,
1983). Reactions: Schaeffer:- ; KOH:- ; Sulphuric
acid: slight; Nitric acid: slight pinkish.
Solitary but abundant, outside the woods,
among grass in lawns, after rains.
Exsiccatum: Pileus light brown with dark
brown fibrils ( 15 J11 [MP]). Lamellae brown with
white efflorescence present. Stipe light brown
270
(10 D6 [MP]) yellowish at the base (11 F6 [MP]).
Material studied: Buenos Aires City, Nufiez,
leg. D. Cabral, O2-V-91 (BAFC: 32.334); Buenos
Aires, Burzaco, leg. E. Albert6é6, O2-X-91 (BAFC:
32.726) and 12-XI-91 (Holotypus: BAFC: 32.765).
Fig. 2: a- Longitudinal section of fruit body. b-
spores and basidium, c- Cheilocystidia, d—- hyphae
of pileus covering.
REMARKS
The specimens studied are similar tO
A.argentinus Speg.(1889) as to pileus and general
aspect , which Spegazzini illustrated with a
photograph in a later work (Spegazzini, 1926),
but they differ in the following features: i-
Spore print. A curious feature is the difficulty
in obtaining one, both in young and mature
specimens of A.pseudoargentinus, whereas in A.
argentinus (Holotypus LPS 15.454 reduced to a
spore print) and later collections by Lindquist,
10-V-1938 (LPS 15.458) and Argentino Martinez,
276
Z0-~LA-T9S9 *CLPS 1574579? the spore-print is
abundant and, furthermore of a very
characteristic reddish ferrugineous colour
(almost as iron oxide); ii- Additionally,
SpegazzZini (op. cit.) indicates for A.argentinus
& double ring and a solid stipe. This contrasts
with the simple ring and fistulose stipe of
A. pseudoargentinus.
Following Heinemann (1962), A. argentinus
pertains to Sect. Agaricus (Campestres). The same
author (pers. comm) believes that A.pseudoargen-—
tinus must tentatively be assigned to Sect.
Sanguinolenti (Mdller et Schaeffer) Singer.
Within Sect. Sanguinolenti, A.pseudoargenti-
nus is close to A.impudicus (Rea) Pilat and
A.lanipes (Médller et Schaeffer) Singer. Agaricus
impudicus differs by its smooth pileus’7 when
young, later, when mature, by its pinkish to dark
brown pileus, with a large and smooth central
disk. Furthermore, it may have membranous
residues of the fugacious general veil, and the
flesh is faintly pinkish at the stem apex. Its
odour is disagreeable, similar to that of Lepiota
cristata. Spores are slightly wider,
cheilocystidia are slightly wider and hyaline.
The species is gregarious and grows in Picea
woods.
Agaricus lanipes, differs by the chocolate
brown cuticule (sometimes tawny or ochraceous) of
this species, which remains for a long time
uncracked and, consequently, does not exhibit
fibrillose squamules from the early stages,
although at later stages it may appear torn into
fibrillose scales and adpressed fibrils and show
the white flesh undernearth (Capelli 1984).
However, the central disk usually remains smooth
or rugose-smooth; the stem is 40-80 mm long and
25-33 mm thick, being rather short and clavate.
According to Miller (1950). the Schaeffer
reaction is usually negative but positive in the
yellowish areas of the stem. The flesh is whitish
varying to “apricot yellow" towards the base and
sometimes yellowish red towards the apex. Its
277
spores are slightly larger in both dimensions. It
grows in broadleaved woods, mainly of Quercus
ilex.
Agaricus fuscofibrillosus differs by its
hazel-brown cap cuticle, densely fibrillose
except at the disc; the stem is relatively
slender, almost perfectly cylindrical and flesh
turning red on cutting. The ring is simple but
narrow and fragile. A. fuscofibrillosus is a wood
inhabiting-species, Capelli,( op cit.) only found
it in the Boscone della Mesola under Quercus
ilex.
ACKNOWLEDGMENTS
We are very grateful to Dr. Paul Heinemann
for his revision of the herbarium samples. We
thank also Dr. D. Cabral for providing fresh
material of Agaricus spp., and the CONICET for
financial support for this project.
Drs P. Heinemann (Gembloux) and Roy Halling
(NY) read the original typescript and made
invaluable suggestions for which we thank them
most heartily. We also thank Mons. J. P. Macarone
for the Latin revision.
LITERATURE .
Capelli, A.1984. Agaricus. Saronno, Italia, 560p.
Heinemann, P. 1962. Agarici Austroamericani V.
Etude des types de C. Spegazzini. Bull. Rech.
Inst. Agron. et Stat. Rech. Gembloux 30: 273-
2382.
Heinemann, P. 1983. Clé de determination de
Micropsalliota (Agaricaceae) et description de
deux espéces nouvelles. Bull. Jard. Bot. Nat.
Belg. 53: 85-95.
Heinemann, P. 1986. Agarici Austroamericani VI.
Apercu sur les Agaricus de Patagonie et de la
278
Terre de Feu. Bull. Jard. Bot. Nat. Belg. 56:
417-446.
Heinemann, P. 1987. Clave para la determinacién
de las especies de Agaricus (Agaricales) de la
Patagonia y de Tierra del Fuego. Darwiniana
28(1-4): 283-291.
Heinemann, P. 1990. Agarici Austroamericani VII.
Agaricaceae des zones tempérées de 1l°Argentine
et du Chili. Bull. Jard. Bot. Nat. Belg. 60:
332-370!
Maerz A. and Paul M., 1930. Dictionary of color.
Mc. Graw Hill Book Company, Inc. New York.
2077p.
Martinez, A. 1949. Agaricaceas nuevas para la
Argentina. Lilloa 21: 43-52.
Martinez, A. 1957. Nueva sp. del género Agaricus.
Rev. Inv. Agri. Bs.As. 11. 3. 238-330.
Miller,F. 1952. Danish Psalliota
species:preliminary studies for a monograph on
the Danish Psalliotae. Part.II. Friesia. 4(3),
204.
Munsell, 1954. Determination of soil color.U.S.
Dep. Agriculture Handbook. Munsell Color
Company, Inc. 16p.
Raithelhuber, J. 1977. Hongos Argentinos II. Bs.
As, 1939p.
Raithelhuber, J. 1988. Flora Mycologica
Argentina. Hongos II. Stuttgart. 287p.
Spegazzini, C. 1899. Fungi Argentini novi vel
critici. Anales Mus. Nat. Bs. As. VI: 141-147.
Spegazzini, C. 1926. Micologia Argentina. Bol.
Acad. Nac. Cienc. Cordoba, 28:325-332.
MY COTAXON
Volume L, pp. 279-289 January-March 1994
THREE NEW SOUTHAMERICAN SPECIES OF BOVISTA
(GASTEROMYCETES)
V. L. Suarez and J. E. Wright
Departamento de Ciencias Bioldgicas
Facultad de Ciencias Exactas y Naturales
Universidad de Buenos Aires
1428 Buenos Aires, Argentina
Summary
Bovista fuegiana, B. singeri and B. sublaevispora are
reported as new species from Southern South America, re-
corded from Tierra del Fuego, Argentina, Nor-—Yungas, Boli-
via and Viffa del Mar, Chile, respectively.
During a survey of the Austroamerican Lycoperdales, se-
veral collections pertaining to the genus Bovista Pers.:
Pers. that did not match any of the described species (cfr.
Coker & Couch, 1928; Wright, 1949; Smarda, 1958; Kreisel,
1967; Calonge & Demoulin, 1975; Jiilich, 1984; Ortega &
Buendia, 1985, 1989) were studied.
For this reason we have decided to propose them as new
species.
Materials and methods
For the study of the fruitbodies of these species the
procedure followed is identical to that given earlier
(Wright & Suarez, 1990). We added two methods, suggested
by Drs. V. Demoulin and 0. K. Miller, respectively:1) soak
ing the material in ETOH 95%, then water, and mounting in
3% KOH + Congo Red, and 2) boiling the sections (under the
cover slip) in chloral hydrate (twice the amount of chlor-
al to that of water). Colours are annotated according to
Maerz & Paul (1930), indicated as MP. Herbarium abbreviat-
ions are according to Holmgren, Holmgren & Barnett (1991).
Bovista fuegiana Suarez & Wright n. sp. (Figs. 1-4; 13)
Gastrocarpo globoso, 2./7-3,cm diam., brunneo, rhizomor-
pho absens. Peridio fragile, in duos stratos: exoperidio
brunneo, laevi, furfuraceo, endoperidio spadiceo. Dehiscen
tia ignota. Gleba brunneo-viridi. pulveracea. Subgleba com
280
pacta, 4-5 mm alt.
Exoperidio pseudoparenchymatico, cellulis 10-31 x 16-50
um. Endoperidio ab hyphis parallelis, juxtapositis, indis-—
tinctis. Sporis globosis pallide brunneis, valde verruco-—
Sis sub lente; sub microscopio electronico dense verruco-—
SiS; “VErerucis rotundatis majores, ev minores, 5.9-6. 9 0m.
Capillitio transitionis, pallide brunneo, poris numerosis
usque ad 1 um diam., sine trabes, ramis principalis 5-8
uum diam.
In pratis sub Berberis prope sylvam. Holotypus: Argenti
na, Fuegiae, Ushuaia, leg. Wright & Del Busto n° 2180, cra
Il .1973;, “in “Herbs: BAFC m3xs2210vconservatus rest.
Fruitbody globose (Fig. 1), 27-30 mm diam., brown, with
out a rhizomorph. Peridium fragile, formed by a brown exo-
peridium (ca MP Pl 8 11-12 J-L), smooth-furfuraceous, and
a "beige" endoperidium (ca MP Pl 14 8 E-F). Dehiscence not
observed due to the material being torn. Gleba greenish-
brown, powdery. Subgleba compact, 4-5 mm high.
Endoperidium (Fig. 2) formed mainly by pseudoparenchy-
matic cells, 10-31 x 16-50 um diam. Endoperidium formed
by juxtaposed parallel hyphae, difficult to measure. Spor-
es globose, light brown, strongly ornamented, appearing
verrucose and without pedicels under LM (Fig. 4); under
SEM the ornamentation is seen as densely arranged, capita-
te warts and among them, much smaller ones; with conspic-
uous pedicels (Fig. 13); 5.5-6.5 um diam. Capillitium (Fig
3-4) of the "transition" type, light brown, with numerous
pores, distributed in principal branches and intermediate
branches, up to 1 um diam., without septa; principal bran-
ches 5-8 um diam., intermediate ones 3-4.8 wm, tapering
to a fine point less than 1-1.8 um diam.
Habitat: in prairies with Berberis, just outside the
forest.
Distribution: Argentina, Tierra del Fuego.
Remarks: The present species is characterized by hav-
ing a "transition" type capillitium with pores, and very
ornamented spores, without pedicels, under LM. It is close
to Bovista aestivalis (Bon.) var. perverrucispora Ortega
& Buendia (Ortega & Buendia, 1989), but differs in the
smaller size of the spores (3.5-4.2 um diam.),
the less crowded ornamentation under SEM of the latter,
and also with conspicuous pedicels. Kreisel (1967) records
B. aestivalis under B. polymorpha (Vitt.)Kreisel. In this
author's key to the species of Bovista, our species would
key out as a member of subgenus Globaria, Ser. Polymorphae
281
Figs. 1-4. Bovista fuegiana Suarez & Wright n.sp. 1: Gene-
ral aspect of gastrocarp, ca. 1 x. 2: Pseudoparenchymatic
cells of the exoperidium. 3: Transition-type of capillit-—
ium. 4: Spores and capillitium under LM.
leading through the features of its exoperidium and the
presence of a gleba to B. aenea Kreisel (UPS!) which, how-
ever, has sinaller spores (3.5-4 um) and exoperidium (15-27
x 11-19 um).
According to V. Demoulin (pers. comm.) it might also
be close to B. cunninghami Kreisel (Kreisel, 1967) (PDD!),
which was described from Australia on the basis of a sin-
gle fruitbody. This appears to be a Species with imperfect-
ly known variability. B. fuegiana differs from it, however,
282
by having: 1) a distinct sterile base, ii) grossly ornamen
ted spores, iii) a much larger fruitbody, and iv) a brown
furfuraceous exoperidium, whereas B. cunninghaini was descr
ibed as lacking a sterile base, having a pseudorrhiza,
smooth spores, a fruitbody of 13 cm diam. and a grumous
whitish exoperidium.
Bovista singeri Suarez & Wright n. sp. (Figs. 5!8; 14)
Gastrocarpo parvo, 8-13 mm diam., globoso-depresso,
brunneo, absque rhizomorpho vel parve evoluto. Exoperidio
tenui, ephemero, verrucis minutis fuscatibus exornato. En-
doperidio tenui, papyraceo, laevi, sapiceo, ab pori apica-
libus regulari dehiscente. Gleba brunneo viridi, pulvera-—
ceacea. Subgleba absens.
Exoperidio vesiculoso, sphaerocystis 12-27 x 16-41 wm.
Endoperidio membranaceo, ab hyphis curtis tenuitunicatis
hyalinis efformantibus, 1.8-5.4 um diam., aliquot pseudo-
parietibus exornatis, alieae ramificationes dichotomicas
efformantibus. Sporis ovoideis, pallide brunneis, verruco-
Sis sub lente, uniguttulatis, apedicellatis vel mucronatis
Sub microscopio electronico verrucis cylindricis, aggrega-
tis, confluentibus vel anastomosantibus. Capillitio typo
Lycoperdon, pallide brunneo, poris .32 wm diam., sine tra-
bes, ramificatione dichotomicis, 1-5 wm diam. Holotypus:
ad foliiis putrefactis®™ in sylvis, alt. 1450/m, SBOLIVIA,;
Dept? La Paz, Prov. Nor-—Yungas, Charobamba, leg. R. Singer
n°? 1129, 13. DD.1996 fein Herb. BARC-on?: S14) 1" conservacus
est. Ad Cl.emycologo RolfSinger dacatus*est.
Fruitbody. small (Fig. 5), 8-13 mm diam., globose-de
pressed, brown, rhizomorph absent or hardly developed. Exo
peridium thin, ephemerous, with minute dark verrucae. Endo
peridium thin, papyraceous, smooth, "beige" (ca MP pl 13,
6 C-E; pl 14, 7 E-F). Dehiscence by a small, irregular,
apical pore. Gleba olivaceous "beige", powdery. Subgleba
absent.
Exoperidium (Fig. 6) vesiculose, formed by sphaerocysts
12-27 x 16-41 wm. Endoperidium membranous, formed by short
thin-walled, hyaline hyphae, 1.8-5.4 um diam., a few with
Figs. 5-8. Bovista singeri Suarez & Wright n. sp. 5: Gene-
ral aspect of gastrocarp, ca. 1 x. 6: long. section of the
peridium with the pseudoparenchymatic cells and the hyphal
structure of the endoperidium. 7: Lycoperdon-type of capi-
llitium. 8: Spores and capillitium under LM.
Oe
9,
eyed
ae
PON
SS
os
ve
ae
4
:
284
pseudowalls and a few others with dichotonious branching.
Spores ovoid, never globose, brown, strongly ornamented;
under LM they appear verrucose, uniguttulate, without pedi
cels,/iwith> very. short mucros (Fig. 8)3°3.2-3.6 xs226<402
um; under SEM the ornamentation appears as somewhat crow-
ded, cylindric verrucae which are nearly free or confluent
united by low connective threads; a short, broken pedicel
is obvious (Fig. 14). Capillitium (Figs. 7-8) of the Lyco-
perdon-type, light brown, with pores ca .3 wm diam., uni-
Formly distributed along the branches, without septa, bran
ching dichotomous, Y-shaped, fairly flexuous, uniform Jn
size, 1-5 wm diam., extremities 1-1.4 um diam.
Habitat: on plant debris in the rain forest, alt. 145C
m.
Distribution: Bolivia.
Material studied: The holotype.
Remarks: This 7s characterized by having a Lycoperdon-
type capillitium with pores, and very ornamented, ovoic
spores without pedicels under LM. It is close to B. gla-
cialis Kreisel (M!), B. dermoxantha (Vitt.)De Toni, and
B. delicata Berk. & Curt. (PCT); the former two, however,
have globose spores, although also strongly ornamented;
the latter has also globose though hardly ornamented spor-
es. Unfortunately, the collector did not give more accur-
ate information concerning the locality or the ecology of
the species.
According to V. Dewoulin (pers. comm.), B. singeri
might be close to B. oblongispora Villiers et al. (1989),
however distinct it may be. We beTieve the differences bet
ween these two species are so obvious to warrant their sep
aration, for B. oblongisporum has: i) a conspicuous cord
of mycelial threads, 77) a capillitium of the "“intermedia-
te" type, iii) larger spores, with different ornamentation
under SEM and iv) generally long spore-pedicels, none of
which features can be found in B. singeri.
Bovista sublaevispora Suarez & Wright n. Sp.
(Bigs '9=12 301s)
Gastrocarpo subgloboso, ca 30 mm diam., brunneo-aeneo
minute punctato, rhizomorpho bene evoluto. Peridio tenui,
Figs. 9-12. Bovista sublaevispora Suarez & Wright n.
sp. 9:, General -aspect of .gastrocarp 7'ca 1 x.) 303) long.
section of the peridium with the pseudoparenchymatic cells
of the exoperidium and the compact hyphal structure of the
endoperidium, 11: Bovista-type of capillitium. 12: Spores
and capillitium under LM.
285
286
15
Figs. 13-15. Photomicrographs of spores under SEM. 13: Bo-
vista fuegiana (Holotype). 14: 8. singeri (Holotype). 15:
B. sublaevispora (Holotype).
papyraceo, duplo. Exoperidio tenui, verrucoso, verrucis
rotundatis vel conicis, fuscis, dense aggregatis. Endope-
ridio papyraceo, aeneo, ab poro apicali irregularibus de-
hiscens. Gleba pallide brunnea vel flavae,pulverulenta.
Subgleba minuta, compacta. Endoperidio pseudoparenchymati-
co, cellulis verrucis 9-29 x 13-43 wm, basis strato amor-
oho vel pséeudoparenchymatico fatiscentibus. Endoperidio
papyraceo, hyphis parallelis juxtapositis instructis, sub-
ter hyphis hyalinis tenuitunicatis, 1.5-4 um diam. Sporis
globosis vel subglobosis, apedicellatis, sub microscopio
287
electronico verrucis minutis, rotundatis, 3.4-5 wm diam.
Capillitio typo Bovista, brunneo, sine poris, trabes ab-
sens, ramis dichotomicis, ramis principalis 7-32 mum diam.
Holotypus: CHILE, Vina del Mar, Horto Botanico, leg. anon.
28.11.1966, in Herb. BAFC n° 32211 conservatus est.
Fruitbody subglobose (Fig. 9), ca 30 mm diam., light
brilliant bronze, with minute dark and densely arranged
punctuations; rhizomorph well developed. Peridium thin,
papyraceous. Exoperidium, thin, under LM blunt to conic,
dark, densely arranged verrucae can be observed. Endoperi-
dium papyraceous, bronze-coloured (ca MP pl 15, 9-10 C-E),
smooth. Dehiscence through an irregular apical aperture.
Gleba light brown to yellowish, powdery. Subgleba very
small, compact, probably not sufficiently mature.
Exoperidium entirely pseudoparenchymatic, the verrucae
formed by cells ca 9-29 x 13-43 wm; the base is an amorph-
ous layer which in some sections suggests having also been
formed by pseudoparenchymatic cells which have disintegra-
ted (Fig. 10)). Endoperidium (Fig. 10) papyraceous, formed
by juxtaposed parallel hyphae which are difficult to meas-
ure, followed by thin-walled hyaline hyphae, some with fal
se septa, 1.5-4 um diam. Spores almost smooth, uniguttula-
te, without pedicels (Fig. 12); under SEM the ornamentat-
ion appears as incipient blunt verrucae not densely arran-
ged and without any sort of connections between them (Fig.
15), 3.5-5 um diam. Capillitium (Figs. 11-12) Bovista-type
brown, without pores nor septa nor pseudosepta, branching
dichotomous, principal branches ca 7-3] wm diam., termin-
al branches ca 1-1.5(-2) ym, the units being flexuous,
each exceeding a microscopic field.
Habitat: no data.
Distribution: Chile.
Material studied: the holotype.
Remarks; This species is characterized by having a Bo-
vista-type capillitium without pores, and globose, almost
smooth spores, which appear slightly verrucose under SEM
and without pedicels. It is close to B. fulva Mass. (K!),
which has subglobose spores but with Tong pedicels. The
latter species belongs to the group of species remaining
attached to the substratum (Subgenus Bovista, Sect. Nanno-
bovista, Ser. Fuscae, according to Kreisel (1967), but we
have no information concerning this feature in our species
other than the presence of a conspicuous rhizomorph at its
base. Furthermore, the new species does not have a "furfu-
raceous-areolate" exoperidium, and its cells are much sma-
ller than in B. fulva (4-7.5 wm diam.).
288
According to V. Demoulin (pers. comm.), B. sublaevispo-
ra might be mistaken with B. coprophila (Cke. & Mass.)G.
H. Cunningh. (K!), but the spore ornamentation of both
Species under SEM is entirely different. Furthermore, the
diameter of the capillitium in B. sublaevispora is wider
(31 um) than that recorded for B. coprophila (23 um).
Acknowledgements
We wish to thank Drs. Orson K. Miller (Virginia Poly-
technic Institute) and Vincent Demoulin (Liége), for cri-
tically reading the typescript and making many valuable
Suggestions.
Keepers of the K, M, PC, UPS, PDD and LPS herbaria are
gratefully acknowledged for the loan of type materials in
their keeping.
Special thanks are due Dr. Gabriel Moreno, Universidad
de Alcala de Henares (Spain), and his team, for invaluable
aid with some of the SEM micrographs.
Literature
Calonge, F. D. & V. Demoulin. 1975. Les Gasteromycétes
d'Espagne. Bull. Soc. Mycol. France 91: 247-292
Coker, W. C. & J. N. Couch. 1928. The Gasteromycetes
of ‘Eastern U. S. and Canada. Chapel Hill, University of
North Carolina Press.
Holmgren, P. K. N. H. Holmgren & L. C. Barnett. 1990.
index: ‘Herbariorum, “Sthered.,. 693.0. Int. Ass. Pin) Laxon:
New York Botanical Garden.
Julich, W. 1984. Die Nichblatterpilze, Gallertpilze und
Bauchpilze. Kleine Kryptogamonflora, Band II, b/1. Gustav
Fisher Verlag. Stuttgart—New York.
Kreisel, HH. 1967. Taxonomisch-—Pf lan Zzengeogma—
phische Monographie der Gattung. Bovista. Beih.
Nowa HedWwigianeoe.244 Pp. fofig. Cramer wee tive:
Maerz, A. & M. R.~ Paul. 1930. A Dictionary of Color.
McGraw-Hill, New York.
Ortega, A. & A. Buendia. 1985. Estudio de algunas espe-
cies con esporas oblongas del género Bovista. Pers. Crypto
gamie, Mycol. 6: 281-288.
Ortega, A. & A. Buendia. 1989. Estudio del complejo Bo-
vista aestivalis (Bon.)Demoulin-B. pusilla (Batsch)Pers.
sensu Kreisel en Espana. Cryptogamie, Mycol. 10 (1): 9-18.
Smarda, F. 1958. Lycoperdaceae in Pilat, A. (Ed.) Flo-
ra’ CSR, Beco oort Oot? a Praia
Villiers jr., J. de, A. Eicker & G. C. A. van der West—
huizen. 1989. Notes on the structure of Bovista oblongispo
289
ra and amplified description of the species. S. Afr.
Tydskr. Plantk. 55 (2): 154-155.
Wright, J. E. 1949. Contribucién al Catalogo de Gastero
myCetes argentinos. 1. Lilloa!242) 191-224.
Wright, J. E. & V. L. Suarez. 1990. South American Gas_—
teromycetes. IV. The genus Abstoma. CUVOce BOt. Te 372-383.
MY COTAXON
Volume L, pp. 291-299 January-March 1994
FOUR NEW SPECIES IN THE LICHEN GENUS PARMELIA
(ASCOMYCOTINA, LECANORALES) FROM SOUTHERN
AFRICA, WITH NOTES ON SOUTHERN AFRICAN
LICHENS *
FRANKLIN A. BRUSSE
National Botanical Institute,
Private Bag X101, Pretoria.
OOO! SOUTH AFRICA.
ABSTRACT
Four new species in the lichen genus Parmelia (Parmeliaceae,
Ascomycotina) are described from southern Africa. They are: Parmelia
chionophila Brusse, P. emolumenta Brusse, P. mesmerizans Brusse and P.
supposita Brusse. One new combination is made: Parmelia protoquintaria
(Hale) Brusse. Notes on several species are provided.
NEW SPECIES
Parmelia chionophila Brusse, sp. nov. Fig3i3?
Thallus foliosus, saxicolus, usque ad 7 cm diam., arcte adnatus. Lobi
elongati, 0.5 - 2.5 mm lati, 100 - 350 um crassi. Thallus superne flavo-viridis,
hebetatus, isidiis sorediisque destitutus; epicortice rudimentali. Cortex superior
9 - 20 um crassus. Stratum gonidiale 30 - 90 um crassum, algis Trebouxiis.
WetiP albida, 50 - 230 um crassa. Cortex inferior 6 - 12 um crassus. Thallus
inferne pallide brunneus, sat rhizinatus. Rhizinae simplices, parvae, ad 0.3 mm
longae, prope basin 25 - 70 um crassae. Apothecia adnata, usque ad 2 mm
diam., act atris, subnitidis. Hypothecium hyalinum, 30 - 40 um crassum, J-.
Subhymenium hyalinum, 20 - 30 um crassum, J+ pallide caeruleum.
Pet Meare bani badium, N+ fugace cyaneum. Hymenium hyalinum, 50 -
60 wm altum, J+ caeruleum. Asci clavati, cum tholis 14 caeruleis (fig. 1).
Ascosporae non visae. Pycnidia hyalina, globosa, circa 100 um diam.
Spermatia acicularia, hyalina, recta, 7.5 - 15.5 X 0.7 um. Thallus acidum
usnicum, acidum hyposticticum, acidum hyposalazinicum, et acidum
hypoconsticticum continens.
TYPUS: SOUTH AFRICA, CAPE PROVINCE - 3319 (Worcester):
* This contribution is dedicated in memory of the late Dr. Ove
Almborn (Lund), an enthusiastic student of southern African lichens,
in recognition of his work on foliose and fruticose Teloschistaceae,
At in gratitude for all his efforts towards a Lichen Flora of southern
rica.
292
-BC (De Doorns) Hex River Mountains. Summit of Matroos Mountain, near
De Doorns. On SE sloped face of Table Mountain Sandstone outcrop on SE
slope with no cliffs nearby. Hex River Valley clearly visible from here. Alt.
ee 0 a F. Brusse 6166, 28. xi. 1991 (PRE, holo-; BM, LD, UPS, iso-).
igura 3.
Thallus foliose, saxicolous, up to 7 cm across, tightly adnate. Lobes
elongate, 0.5 - 2.5 mm broad, 100 - 350 um thick. Upper surface yellow-
presi matt, neither isidiate nor sorediate; epicortex rudimentary. Upper cortex
- 20 um thick. Algal layer 30 - 90 um thick, algae Trebouxia. Medulla
whitish, 50 - 230 um thick. Lower cortex 6 - 12 um thick. Lower surface pale
brown, darkening towards the lobe tips and margins, moderately rhizinate.
Rhizines simple, small, to 0.3 mm long, 25 - 70 um thick near base. Apothecia
adnate, up to 2 mm across, disc black, subglossy. Hypothecium hyaline, 30 -
40 um thick, J-. Subhymenium hyaline, 20 - 30 um thick, J+ ane blue.
Epihymenium toffee brown, N+ fleeting blue. Hymenium hyaline, 50 - 60 um
high, J+ blue. Asci clavate; tholus J+ Flue (fig. 1). Ascospores not seen.
Pycnidia hyaline, globose, ca. 100 um diam. Spermatia hyaline, straight,
needles, 7.5 - 15.5 X 0.7 um. Chemistry: usnic, hypostictic, hyposalazinic, and
hypoconstictic acids present.
Etymology: chion (G) = snow; philos (G) = loving; alluding to the relatively
large amount of snow that falls at the type locality. Due to this fact, this is a
favourite spot for local skiers.
his new species resembles the Australian Parmelia subcrustacea
Gyeln. (Elix 1981), but is clearly distinguished from it by the N+ fleeting blue
epihymenium, which is in contrast to the N- epihymenium of the latter.
Parmelia chionophila also becomes coarse-pruinose towards the tips of the
lobes, and lacks a coherent epicortex. P. subcrustacea, on the other hand,
becomes more glossy towards the lobe tips and possesses a pored epicortex.
The N+ fleeting blue epihymenial reaction which was discussed in detail by
Brusse (1993), is present in many effigurate-crustose brown Parmeliae and in
several lichens assigned to Karoowia by Hale (1989b), including the type, P.
adhaerens Ny1. P. chionophila is the largest and most foliose lichen found so
far, with the N+ fleeting blue epihymenial reaction.
Thus far this new species has been discovered only at the type
locality, Matroos Mountain the highest peak in the Hex River Mountains, near
De Doorns.
Parmelia emolumenta Brusse, sp. nov. Fig. 4.
Thallus foliosus, saxicolus, usque ad 5 cm diam., arctissime adnatus.
Lobi elongati, 0.2 - 1.5 mm lati, 150 - 700 um crassi. Thallus superne flavo-
viridis, hebetatus vel apicem loborum versus grosse pruinosus, isidiis
sorediisque destitutus; epicortice rudimentali. Cortex superior circa 20 wm
crassus. Stratum gonidiale 25 - 140 wm crassum, algis Trebouxiis. Medulla
albida, 70 - 600 um crassa. Cortex inferior 10 - 15 um crassus. Thallus inferne
pallide brunneus. Rhizinae non bene evolutae. Apothecia (immatura tantum
visa) ad 1 mm diam., adnata. Epihymenium pallide badium, N+ fugace ;
cyaneum. Pycnidia ae globosa, circa 120 wm diam. Spermatia acicularia,
hyalina, recta, 5.5 - 8.0 X 0.7 wm. Thallus acidum usnicum, acidum sticticum,
acidum consticticum et acidum norsticticum (min.) continens.
TYPUS: SOUTH AFRICA, CAPE PROVINCE - 3319 (Worcester):
-BC (De Doorns) Hex River Mountains. Summit of Matroos Mountain, near
293
FIGURE 1. - Parmelia chionophila Brusse, ascus and paraphyses. F. Brusse
6766, holotype. Bar = 10 ym.
FIGURE 2. - Parmelia supposita Brusse, ascus and paraphyses. F. Brusse
5496, holotype. Bar = 10 wm.
De Doorns. On SE sloped face of Table Mountain Sandstone outcrop on SE
slope with no cliffs nearby. Hex River Valley clearly visible from here. Alt.
2240 m. F. Brusse 6165, 28. xi. 1991 (PRE, holo-; LD, iso-). Figura 4.
Thallus foliose, saxicolous, up to 5 cm across, very tightly adnate.
Lobes elongate, 0.2 - 1.5 mm broad, 150 - 700 um thick. Upper surface
yellow-green, matt to coarsely pruinose towards the lobe tips, both isidia and
soredia lacking; epicortex enh mente Upper cortex ca. 20 um thick. Algal
layer 25 - 140 um thick, algae Trebouxia. Medulla whitish, 70 - 600 um thick.
Lower cortex 10 - 15 wm thick. Lower surface pale brown, darkening towards
the tips and margins. Rhizines poorly developed. Apothecia (only immatures
seen) up to | mm across, adnate. Epihymenium fetes brown, N+ fleeting
blue. Pycnidia hyaline, globose, ca. 120 wm diam. Spermatia hyaline, straight
needles, 5.5 - 8.0 X 0.7 wm. Chemistry: usnic, stictic, constictic acids, and
norstictic (min.) acids present.
Parmelia emolumenta is another new species with an N+ fleeting
blue epihymenium. It otherwise seems similar to Xanthoparmelia
xanthomelanoides Elix & Johnston a species with an N- epihymenium, and also
a glossy pored-epicorticate upper surface. P. emolumenta lacks a coherent
epicortex.
This new species is so far known only from the type locality,
Matroos Mountain, near De Doorns.
294
Parmelia mesmerizans Brusse, sp. nov. Fig).
Thallus foliosus, saxicolus, usque ad 7 cm diam., arctissime adnatus.
Lobi elongati, 0.3 - 2.0 mm lati, 130 - 380 um crassi. Thallus superne flavo-
viridis, hebetatus vel apicem loborum versus scabrosus, isidiis sorediisque
destitutus; epicortice rudimentali. Cortex superior circa 20 um crassus. Stratum
gonidiale 30 - 100 um crassum, algis Trebouxiis. Medulla albida, 60 - 270 um
crassa. Cortex inferior 8 - 12 um crassus. Thallus inferne atro-brunneus vel
piceus, sat rhizinatus. Rhizinae simplices, parvae, usque ad 150 um longae,
prope basin 45 - 150 um crassae. Apothecia non visa. Pycnidia hyalina,
lobosa, circa 120 wm diam. Spermatia acicularia, hyalina, recta, 6.5 - 14.5 X
.8 um. Thallus acidum usnicum, acidum hyposticticum, acidum
hyposalazinicum, et acidum hypoconsticticum continens.
TYPUS: SOUTH AFRICA, CAPE PROVINCE - 3319 (Worcester):
-BC (De Doorns) Hex River Mountains. Summit of Matroos Mountain, near
De Doorns. On level Table Mountain Sandstone pavement on SE slope, near
summit. Full sun. Alt. 2230 m. F. Brusse 6/41, BB. xi. 1991 (PRE, holo-;
LD, iso-). Figura 5.
Thallus foliose, saxicolous, up to 7 cm across, very tightly adnate.
Lobes elongate, 0.3 - 2.0 mm broad, 130 - 380 um thick. Upper surface
yellow-green, matt to scabrous towards the lobe tips, non-isidiate and non-
sorediate; epicortex rudimentary. Upper cortex ca. 20 ym thick. Algal layer 30
- 100 pm thick, algae Trebouxia. Media whitish, 60 - 270 um thick. Lower
cortex 8 - 12 um thick. Lower surface dark brown to black, becoming lighter
brown towards the lobe tips. Rhizines simple, small, up to 150 pm long, 45 -
150 um thick near base. Apothecia not seen. Pycnidia hyaline, globose, ca. 120
um diam. Spermatia hyaline, straight needles, 6.5 - 14.5 X 0.8 um. Chemistry:
usnic, hypostictic, hyposalazinic, and hypoconstictic acids present.
This new species is closest to Parmelia protoquintaria (Hale)
Brusse, with a similar chemistry and lower surface colour. However, P.
rotoquintaria is most closely related to Parmelia xanthomelaena Mill. Arg.,
bile contains the hypostictic acid cohort instead of the stictic acid series of the
latter. The lobes are sublinear and removable with the lower surface intact
throughout the thallus, whereas with P. mesmerizans this is not possible,
because the centre is subareolate, although bullately so. The marginal lobes of
P. mesmerizans are elongate, and do not possess a pored epicortex.
Conversely, the marginal lobes of P. protoquintaria are sublinear, much less
distinctly differentiated from the central lobes, and possess a pored epicortex.
Parmelia tantillum Brusse (1989; syn.: Xanthoparmelia inconspicua
Hale) is another small species containing the hypostictic acid series, but this
species is pale brown below, and also has a glossy upper surface with a pored
epicortex.
Up until now, this new species is known only from the type locality,
Matroos Mountain, the highest peak in the Hex River Mountains, near De
Doorns.
Parmelia supposita Brusse, sp. nov. Fig. 6.
Thallus effigurato-crustosus, saxicolus, usque ad 3 cm diam.
Hypothallus ater. Lobi sublineares, usque ad 3 mm longi, 0.1 - 0.5 mm lati, 70
- 80 um crassi. Thallus superne flavo-viridis, subnitidus, isidiis sorediisque
destitutus, epicortice destituto. Cortex superior 15 - 20 um crassus. Stratum
20
gonidiale 20 - 50 um crassum, algis Trebouxiis. Medulla albida, 10 - 110 um
crassa. Cortex inferior 7-9 um crassus. Thallus inferne piceus, cum
hypothallo atro bene evolutus. Rhizinae non evolutae. Apothecia immersa vel
sessilia, usque ad 0.4 mm diam., discis badiis vel pallide badiis, saepe
concavis. Hypothecium hyalinum, 15 - 30 um crassum, J-. Subhymenium
hyalinum, circa 10 um crassum, J+ caeruleum. Epihymenium pallide badium,
-. Hymenium hyalinum, 35 - 45 ym altum, J+ caeruleum. Asci clavati, cum
tholis J+ caeruleis (fig. 2). Ascosporae senae vel octonae, hyalinae, simplices,
ellipsoideae, 9.5 - 12 X 5.0 - 6.5 um. Pycnidia eae globosa, 80 - 100 um
diam. Spermatia acicularia, hyalina, recta, 7 - 9.5 X 0.8 um. Thallus acidum
usnicum, acidum hyposticticum, acidum sticticum, acidum hypoconsticticum et
acidum consticticum continens.
TYPUS: SOUTH AFRICA, CAPE PROVINCE - 3319 (Worcester):
-AC (Tulbagh) Obiqua Mountains. Nuwekloof near Tulbagh. Farm: Nieuw
Kloof. On W facing Table Mountain Sandstone kranz, on SW slope, in poort.
at 140 m. F. Brusse 5496, 22. ii. 1988 (PRE, holo-; BM, LD, iso-). Figura
Thallus effigurate-crustose, saxicolous, up to 3 cm across.
Hypothallus black. hGBes sublinear, up to 3 mm long, 0.1 - 0.5 mm wide, 70 -
180 um thick. Upper surface yellow-green, subglossy, not isidiate and not
sorediate; epicortex absent. ove cortex 15 - 20 um thick. abe layer 20 - 50
pm thick, algae Trebouxia. Medulla whitish, 10 - 110 um thick. Lower cortex 7
- 9 wm thick. Lower surface black, with well developed black hypothallus
(sometimes thicker than lobes). Rhizines not developed. Apothecia immersed to
sessile, up to 0.4 mm across, disc chestnut brown or lighter, often concave.
Hypothecium hyaline, 15 - 30 um thick, J-. Subhymenium hyaline, ca. 10 wm
thick, J+ blue. Epihymenium toffee brown, N-. Hymenium Halong 35 - 45 um
high, J+ blue. Asci clavate, six- to Te RHE, tholus J+ blue (fig. 2).
Ascospores hyaline, monolocular, ellipsoid, 9.5 - 12 X 5.0 - 6.5 wm. Pycnidia
hyaline, globose, 80 - 100 um diam. Spermatia hyaline, straight, needles, 7 -
9.5 X 0.8 pum. Chemistry: usnic, hypostictic, stictic, hypoconstictic and
constictic acids present.
This is a very peculiar tiny new species in the artificial section
Xanthoparmelia, in that the lobes are embedded in a black hypothallus, which
is visible as black margins to the lobes under a cea microscope. The
hypothallus can become very thick below, thicker than the thallus sometimes,
and often includes some substrate grains. The hypothallus hyphae arise from
the black lower cortex, which seems to be present throughout the thallus. This
causes the lobes to be more easily removed than expected for an effigurate-
crustose lichen. This development of a hypothallus is unknown in Parmelia,
and consequently Parmelia supposita is a very distinct species. This new
species was at fs assumed to be Parmelia adhaerens Nyl.. with the identical
chemistry, but P. adhaerens is N+ Ree blue in the epihymenium, whereas
Neg SePROSND is not. Although Brusse (1993) thought that all the type material of
Nig aerens is sterile, on checking the records it was found that one piece
(BM) is fertile. This specimen has been re-examined and found to be N+
Tea blue in the epihymenium. This fertile specimen at BM should be
viewed as the holotype of P. adhaerens Nyl., because the material at H-Ny]l is
sterile and only a tiny fragment, and could at best be deemed a Let ae
A hypothallus is also lacking in P. adhaerens, which is attached to
the rock substrate by means of small poorly developed rhizines and, in the
interior, directly to the lower cortex.
The back margined lobes of this novel species, bring to mind other
296
strikingly black margined species such as Parmelia endochromatica (Hale)
Brusse, and Parmelia ponderosa Brusse, but these are larger species with
gyrophoric acid and "schenckiana" pigments in the medulla. These latter
species also do not possess a hypothallus.
This new species has been collected from two localities, Nuwekloof,
just west of the historic town of Tulbagh, and Pakhuis Pass, north-east of
Clanwilliam.
Additional Specimen Examined:
SOUTH AFRICA, CAPE PROVINCE - 3218 (Clanwilliam): -BB
(Clanwilliam) 10 km NE of Clanwilliam, Pakhuis Pass, 50 m W of- and
opposite entrance to Klein Kliphuis farm. On large Table Mountain
Sandstone boulder-outcrop, on SW slope, on S faces, near ground. Alt.
600 m. F. Brusse 4939, 11. ii. 1986 (B, BM, COLO, PRE).
NOTES
The following "species" have been found to be based on material
infected with lichen parasites, and are fraught with warts or galls caused by the
parasites. These warts have previously been misinterpreted as dactyls by myself
(Brusse 1989b, 1991), or sometimes “isidia" by Hale (1986, 1987) and even
pustules that become "coarse subsorediate masses" or "coarse soredia" by Hale
(1986, 1989, 1989a). They are Xanthoparmelia areolata Hale, X. coneruptens
Hale, X. eruptens Hale, 4 evernica Hale, X. pustulifera Hale, X. saniensis
Hale. Some so-called species possess rather elongate linear warts, resembling
incipient linear soralia, and warts or galls of this type have been found on
Xanthoparmelia pustulosorediata Hale and X. rubropustulata Hale, and have
been noticed on several other species such as the brown P. parilis Brusse and
the grey P. molybdiza Nyl. When mature these warts become rather hollow
inside and break open, and then resemble pustules. The type material of
Psiloparmelia arhizinosa Hale may also be similarly infected with a parasite,
causing abnormalities that have been interpreted as pustules erupting into
coarse sorediate masses (Hale 1989a, Elix & Nash 1992). It seems that lichens
Blowing in overhang positions are more susceptible to succumb to infection by
ichen parasites.
Regrettably I have made combinations in Parmelia in error for some
of these lichens, but the status as good species, without the lichen parasite
induced changes, is still unclear for some of them, especially since the parasites
can also effect changes in chemistry. Documented examples of how lichen
Gas can cause changes in chemistry are given by Hawksworth, Paterson &
ote (1993), where the chemistries of various lichens are diversified by
infection. In Parmelia the genera Polycoccum and Nesolechia are
overwhelmingly common as parasites. Nesolechia is a parasite that has acquired
this habit from the lichenized condition in the past (i.e. it is a degenerate
lichen), but there are no reports of chemical changes caused by this parasite as
yet. Polycoccum is a fungus with no history of lichenization, that has adapted to
parasitizing lichens presumably from saprophytic or plant/fungal parasitic
ancestors, and the possibility that this parasite could elaborate on the host
chemistry is remote. However, this type of parasite could still induce the host
to change its chemistry under conditions of infection. Nevertheless, the
chemistry is often not altered, and these cases can clearly be withdrawn for the
time being, as is done in some of the entries below.
Parmelia evernica (Hale) Brusse oe
The original material I saw had a few dark tipped isidium-like
structures on it, and this was the basis of the new combination (Brusse 1989a).
297
However, I have recently received further material of this lichen, and it clearly
is Parmelia lecanoracea Mill. Arg. with warts caused by a lichen parasite.
PARMELIA PROTOQUINTARIA (Hale) Brusse, comb. nov.
Basionym: Xanthoparmelia protoquintaria Hale, Mycotaxon 34:
elu EEE
This species is the hypostictic acid series containing counterpart of
Parmelia xanthomelaena Mill. Arg., which produces the stictic acid cohort.
Parmelia rubropustulata (Hale) Brusse
This is another clear-cut case of a lichen parasite, in this case
infecting P. endomiltodes Nyl. The brown P. parilis Brusse has also been seen
infected, producing the same linear wart structures, which burst or break open,
and then look like pustules.
Parmelia saniensis (Hale) Brusse
When making the new combination Parmelia brevilobata (Hale)
Brusse (1991), the structures on this lichen were thought to be dactyls, but
these are plainly the work of a lichen parasite, and thus P. brevilobata is
synonymous with P. saniensis. The minor substance in this lichen, which I
originally thought may be lividic acid, 1s in fact unknown TE-2 (Esslinger
1977), which occurs in the brown P. fissurina Zahlbr., a common lichen at
higher altitudes in the Cape Province. Although quite similar to P.
subeaitacen Gyeln., P. saniensis contains unknown TE-2 and the pigment
skyrin in the medulla near the lower cortex, in addition to the hypostictic,
SES ale hypoconstictic acids and unknown PQ-3 (the hypostictic acid
cohort), which they both contain (Elix 1981; Elix, Johnston & Armstrong
1986; Hale 1990). P. saniensis has a years priority over P. brevilobata, and is
therefore the correct name for this lichen, despite the presence of the parasite.
ACKNOWLEDGEMENTS
I wish to convey my sincere thanks to Prof. D. L. Hawksworth
(London) for very kindly reviewing this paper. The examination of type and
other valuable material ae B, BM, COLO, FH, G, GLAM, H, Lb. TNS,
TRH, TUR, US, VER, W, & ZT, is gratefully acknowledged. The
photography is the generous work of Mrs A. J. Romanowski, to whom I extend
my thanks. The author is grateful for the friendly co-operation from several
magnanimous private landowners who allowed the author to look for lichens on
their properties. The author is especially grateful to the South African Ski Club
(Cape Town) for the kind permission to use their elegant Peak Hut on the
summit of Matroos Mountain, and for permitting access to their leased ski area
on this mountain.
LITERATURE CITED
BRUSSE, F. A. 1989. Two new species of Parmelia (Parmeliaceae,
Lichenes), further new combinations and notes, and additional new
lichen records from southern Africa. Mycotaxon 34: 399 - 406.
BRUSSE, F. A. 1989a Four new species of Parmelia (Lichenes) from
southern Africa, with further notes, new combinations and new lichen
records. Mycotaxon 35: 21 - 34.
BRUSSE, F. A. 1989b. Three new species in the lichen genus Parmelia
(Parmeliaceae, Ascomycotina) from southern Africa, with further notes.
Mycotaxon 36: 305 - 311.
298
BRUSSE, F. A. 1991. Eight new species in the lichen genus Parmelia
(Parmeliaceae, Ascomycotina) from southern Africa, with notes on
southern African lichens. Mycotaxon 40: 377 - 393.
BRUSSE, F. A. 1993. Eight new species in the lichen genus Parmelia
(Ascomycotina, Parmeliaceae) from southern Africa, with notes on
southern African lichens. Mycotaxon 49: | - 17.
ELIX, J. A. 1981. New species of Parmelia subgen. Xanthoparmelia (Lichens)
aoe Australia and New Zealand. Australian Journal of Botany 29: 349 -
ELIX, J. A., JOHNSTON, J. & ARMSTRONG, P. A. 1986. A revision of the
lichen genus Xanthoparmelia in Australasia. Bulletin of the British
Museum (Natural History) Botany series 15: 163 - 362.
ELIX, J. A. & NASH, T. H. 1992. A synopsis of the lichen genus
aaa (Ascomycotina, Parmeliaceae). The Bryologist 95: 377 -
ESSLINGER, T. L. 1977. A chemosystematic revision of the brown
Parmeliae. Journal of the Hattori Botanical Laboratory 42: 1 - 211.
HALE, M. E. 1986. New species of the lichen genus Xanthoparmelia from
utc Africa (Ascomycotina: Parmeliaceae). Mycotaxon 27: 563 -
HALE, M. E. 1987. Additions to the Xanthoparmelia Flora of southern Africa
(Lichenized Ascomycotina: Parmeliaceae). Mycotaxon 29: 251 - 266.
HALE, M. E. 1989. New species in the lichen genus Xanthoparmelia
(Ascomycotina: Parmeliaceae). Mycotaxon 34: 541 - 564.
HALE, M. E. 1989a. A new lichen genus, Psiloparmelia Hale
(Ascomycotina: Parmeliaceae). Mycotaxon 35: 41 - 44.
HALE, M. E. 1989b. A monograph of the lichen genus Karoowia Hale
(Ascomycotina: Parmeliaceae). Mycotaxon 35: 177 - 198.
HALE, M. E. 1990. A synopsis of the lichen genus Xanthoparmelia (Vainio)
Hale (Ascomycotina, Parmeliaceae). Smithsonian Contributions to
Botany 74: | - 250.
HAWKSWORTH, D. L., PATERSON, R. R. M. & VOTE, N. 1993. An
investigation into the occurrence of metabolites in obligately
lichenicolous fungi from thirty genera. Bibliotheca Lichenologica in
press.
FIGURE 3. - Parmelia chionophila Brusse, habit. F. Brusse 6766,
holotype. Scale in mm.
FIGURE 4. - Parmelia emolumenta Brusse, habit. F. Brusse 6765,
holotype. Scale in mm.
FIGURE 5. - Parmelia mesmerizans Brusse, habit. F. Brusse 6147,
holotype. Scale in mm.
FIGURE 6. - Parmelia supposita Brusse, habit. F. Brusse 5496,
holotype. Scale in mm.
299
MYCOTAXON
Volume L, pp: 301-306 January-March 1994
FURIA SHANDONGENSIS
(ZYGOMYCETES: ENTOMOPHTHORALES),
A NEW PATHOGEN OF EARWIGS !
Weimin Wang
Wenhua Lu
Institute of Atomic Energy Use, Shandong Academy of Agricultural
Sciences, Jinan, Shandong 250100, P. R. China
Zengzhi Li 2
Department of Forestry, Anhui Agricultural University
Hefei, Anhui 230036, P. R. China
Abstract: An epizootic of Forficula sp. was recorded in Eastern
China. The causal agent is an entomophthoralean species that differs
from Erynia forficulae and Furia ellisiana, and is described here as
Furia shandongensis, sp. nov., the third pathogen of earwigs. Its
primary conidia are obovoid, 16.8—22.8 X 9.6-12.0 Um (averaging
1 OOP OS m2 L/D 96-273). “Cystidia and thizoidsare asi wide
as conidiophores, and rhizoids have no terminal holdfasts. Resting
spores are 26.4—36.0 Lm in diameter.
Keywords: Forficulidae, Dermaptera, Entomophthorales, Furia
shandongensis
INTRODUCTION
An epizootic of earwig adults and nymphs (Forficulidae) was observed
in suburban Jinan, Shandong Province, Eastern China, in August 1990.
Cadaver density was 1-2 per m2 in their damp, dark habitats under pots
and shading matting, and in cracks of walls and soil. The cadavers were
! Supported by National Natural Science Foundation of China.
2 Author to whom all correspondence should be addressed.
302
attached to substrates with their legs. The anterior and posterior ends of the
cadavers bent upwards (Fig. 1). White hymenia emerged from the
intersegmental membranes of the thorax and abdomen, kept extending,
and finally joined together making the whole cadaver look creamy. The
great number of conidia discharged around the cadavers indicated an
entomophthoralean infection. These conidia were homogeneous, suggest-
ing that only one fungal species was the pathogen of the epizootic.
MATERIALS AND METHODS
Morphological methods of Li and Humber (1984) were used to make
glass slide preparation for microscopic observation on primary and secondary
conidia, hyphal bodies, conidiophores, cystidia, rhizoids, and resting spores.
Except for aceto-orcein used for nuclear staining, lactophenol-aniline blue
was used for other staining and dissecting work.
For isolation and maintenance, potato dextrose agar (PDA), Sabouraud
dextrose agar (SDA), and coagulated egg yolk + Sabouraud maltose agar
(EYSMA) (Soper et al., 1975) were used. Growth rates on the three
media were compared at 24°C.
RESULTS AND DISCUSSION
The microscopic characteristics of the specimens collected agreed with
the description of the genus Furia (Batko) Humber (1989) but were not
attributable to any described species of this genus. The description of this
new species is based on the microscopic observation of the specimens .
FURIA SHANDONGENSIS Wang, Lu & Li, sp. nov. [Figs. 1-10]
CONIDIA PRIMARIA uninucleata, bitunicata, incolorata, obovoidea,
plerumque symmetrica, 16.8-22. 8 X 9.6-12.0 um (med. 19.2 + 1.2 x
10.3-0.9 Um), L/D= 1.6—2.3). CONIDIA SECUNDARIA obovoidea
sed minora, 13.2-16.8 x 9.6-12.0 Um (med. 14.9+0.9 x 10.1 40.8 Lm,
L/D = 1.2-1.7). Capilliconidia desunt. CORPORA HYPHALIA clavata
vel hyphoidea, 7.2-9.6 Um in diam. CONIDIOPHORA bifurcata, 9.6—
12.0 um in diam. CYSTIDIA aseptata, simplicia, basi 9.6-10.8 [fm in
diam., gradatim angustata. RHIZOIDEA monohyphoidea, Y—ramosa ad
apicem, 9.6—12.2 um basi in diam. SRPORAE PERDURANTES sphaeri-
cae, hyalinae, 26.4-36.0 Um in diam. (med. 31.1 + 2.4 Um).
PRIMARY CONIDIA (Figs. 2-3) uninucleate, bitunicate, colorless, ob-
ovoid, with a rounded apex and a rounded papilla, mostly symmetric,
303
FIGURES 1-10. Furia shandongensis. 1. A dead earwig with fungal
hymenia protruding from intersegmental membranes. 2-3. Primary conidia,
x590. 4-6. Secondary conidia, X590. 7. Conidiophores, X590. 8. A cysti-
dium, X250. 9. A rhizoid, X250. 10. Resting spores, x400.
304
16,8-22°81X\9.6-12:0. ami Waveraping, 1999-6 12: x 11013 009 ie
1.6-2.3, n = 100), with cytoplasm containing 1-2 large vacuoles, forcibly
discharged by papillar eversion. SECONDARY CONIDIA (Figs. 4-6)
similar to primary conidia but smaller, 13.2-16.8 x 9.6-12.0 Lum (averaging
14.94 0.9 xX 10.1 £0.8 um, L/D = 1.2-1.7, n = 100). No capilliconidia
formed. HYPHAL BODIES clavate or hypha-like, 7.2-9.6 um wide,
multinucleate. NUCLEI elliptical to subglobose, 5.1-8.0 x 2.5-5.1 [um in
primary conidia. CONIDIOPHORES bifurcately branched (Fig. 7),
determinate, 9.6-12.0 um in diam., usually covering intersegmental mem-
brane of host bodies or forming continuous pale yellowish hymenia.
CYSTIDIA (Fig. 8) emerging through cuticle of dead host before conidio-
phores, aseptate, unbranched, 9.6-10.8 um wide at base, tapering gradually
to a rounded tip 5.0-6.5 Um wide. RHIZOIDS (Fig. 9) hypha-like, Y-
branched, 9.6-12.2 lm wide at base. RESTING SPORES (Fig. 10)
spherical, hyaline, smooth, 26.4-36.0 fm in diam. (averaging 31.1 + 2.4
lum), with bilayered wall 2.6-3.7 um thick.
Holotype: ACAFP 11025; adult of Forficula sp.; deposited at herbarium
of Forest Protection, Department of Forestry, Anhui Agricultural Uni-
versity. Collected by Weimin Wang.
Paratypes: AEU08820-—1 through 5; adults and nymphs of the same host
species; deposited at Mycological Laboratory, Institute of Atomic
Energy Use, Shandong Academy of Agricultural Sciences. Collected by
Weimin Wang.
Type Host: Forficula sp. (Dermaptera: Forficulidae)
Type Locality: East suburbs of Jinan, Shandong; 20 August 1990.
The following characteristics make this fungus attributable to Furia
(Batko) Humber (1989): uninucleate and bitunicate conidia, bifurcately
branched conidiophores, hypha-like cystidia as wide as conidiophores, and
monohyphal rhizoids which are as wide as conidiophores and have no
strongly differentiated holdfast.
There are two earwig-pathogenic species published. The first was
described as Entomophthora forficulae Giard (1889). Its conidia were long
ellipsoid, 20-25 x 6.8 um. The available information was too limited for
Ben-Ze’ev and Kenneth (1982) to assign the species to any subgenus of
the genus Erynia in their neo-Batkoan classification or for Humber (1989)
to place it among the four genera in his revised classification derived from
Erynia Nowakowski emend. Humber & Ben-Ze’ev (1981). The second
species, Furia ellisiana (Ben-Ze’ev) Humber, was originally described as
Empusa forficulae var. major Petch (1944) and redescribed as Erynia ellisiana
305
Ben-Ze’ev (1986) before its transfer to Furia by Humber (1989). Its pri-
mary conidia were obovoid, multivacuolate, 19.8-30.0 x 12.6—20.2 [m.
Rostrup (1893, cited in Petch, 1944) recorded a fungus as Entomophthora
forficulae on earwigs in Denmark which was different, as is Furia shan-
dongensis, from the above two species by the primary conidia 18-21 x 8-
10 Em, with a vacuole at each end. As Ben-Ze’ev noted, it appeared to be
the third species attacking Dermaptera. The new species described here as
F, shandongensis has similar magnitude to that measured by Rostrup and
cited by Petch (1944), and they probably are the same fungus although the
1 or 2 vacuoles of F. shandongensis are mostly central rather than the polar
ones observed by Rostrup.
In the genus Furia, F. crustosa (MacLeod & Tyrrell, 1979) Humber and
F, ithacensis (Kramer, 1981) Humber have measurements close to that of F.
shandongensis but their resting spores have undulate or deeply incised epi-
spores in comparison with the smooth resting spores of F. shandongensis.
The first species is a pathogen of Lepidoptera and the second is pathogenic
to dipterans whereas F. shandongensis is a pathogen of the Dermaptera.
This fungus could grow on PDA and EYSMA, but not on SDA. Its
growth rate on EYSMA was three times faster than on PDA. The primary
conidia produced on EYSMA were similar in shape and size (19.6 X 10.6
lim) to those produced on the host. Resting spores formed on both media
were ca. 27.1 [tm in diameter and had walls ca. 2.9 lm thick.
ACKNOWLEDGMENTS
We thank Dr. R. A. Humber for his critical review of the manuscript,
and for its preparation and final submission for publication.
PITBRAROURE CILED
Ben-Ze’ev, I. 1986. Notes on Entomophthorales (Zygomycotina)
collected by T. Petch. II. Erynia ellisiana sp. nov., non Erynia forficulae
(Giard) comb. nov., pathogens of Forficulidae (Dermaptera). Mycotaxon
27: 263-269.
Ben-Ze’ev, I. and R. G. Kenneth. 1982. Features-criteria of taxonomic
value in the Entomophthorales. II. A revision of the genus Erynia
Nowakowski 1881 (= Zoophthora Batko, 1964)). Mycotaxon 14: 456-
475.
Giard, A. 1889. Sur quelques types remarquables de champignons
entomophytes. Bull. Sci. Fr. & Belg. 20: 197-224.
306
Humber, R. A. 1989. Synopsis of a revised classification for the Ento-
mophthorales (Zygomycotina). Mycotaxon 34: 441-460.
Humber, R. A. and I. Ben-Ze’ev. 1981. Erynia (Zygomycetes: Ento-
mophthorales) emendations, synonymy and transfers. Mycotaxon 13:
506-516.
Kramer, J. P. 1981. A mycosis of the blood-sucking snipe fly,
Symphoromyia hirta, caused by Erynia ithacensis sp. nov. Mycopathologia
75: 159-164.
Li, Z. and R. A. Humber. 1984. Erynia pieris (Zygomycetes:
Entomophthoraceae), a new pathogen of Pieris rapae (Lepidoptera:
Pieridae): description, host range, and notes of Erynia virescens. Can. J.
Bot. 62: 653-663.
MacLeod, D. M. and D. Tyrrell. 1979. Entomophthora crustosa n. sp. as
a pathogen of the forest tent caterpillar, Malacosoma disstria. Can.
Entomol. 111: 1137-1144.
Petch, T. 1944. Notes on entomogenous fungi. Trans. Brit. Mycol. Soc.
27: 81-93.
Remaudiére, G. and S. Keller. 1980. Revision systématique des genres
d’Entomophthoraceae a potentialite entomopathogéne. Mycotaxon 11:
323-338.
MY COTAXON
Volume L, pp. 307-314 January-March 1994
TWO NEW PATHOGENS
OF DIPTERAN INSECTS !
€
Meizhen Fan
Department of Forest Protection, Northwest College of Forestry
Yanlingzhen, Shaanxi 712100, P. R. China
Zengzhi Li 2
Department of Forestry, Anhui Agricultural University
Hefei, Anhui 230036, P. R. China
Abstract: Two new entomophthoralean fungi pathogenic on dip-
teran insects were recorded in China. Pandora shaanxiensis was
found on blue bottle flies in Shaanxi, Northwestern China. Its
primary conidia are obovoid, symmetric, 30.3—43.2 x 12.5-19.8 Um
(averaging 32.7 x 15.0 um), L/D = 1.7-3.2 (averaging 2.4). Cysti-
dia are 2 times the thickness of conidiophores, and rhizoids end in a
discoid holdfast. Erynia chironomis was found in an epizootic of
chironomid midges in Anhui, Eastern China. Its primary conidia are
narrow, obovoid, usually asymmetric and sometimes curved, the
largest diameter in the apical half, 29.260.6 x 10.4-21.9 Um (aver-
acing 64 11—43.0).x115.5-17- 1 mam) 0/1 2.1-4.3 (averagine 2:5—
3.6). Cystidia are 203 time the thickness of conidiophores. Rhizoids
are 2 times the thickness of conidiophores and do not terminate in
terminal discoidal holdfasts.
Keywords: Entomophthorales; Pandora shaanxiensis; Erynia
chironomis, Diptera, Calliphoridae, Chironomidae
' Supported by National Natural Science Foundation of China.
2 Author to whom all correspondence should be addressed.
308
Insects of the Diptera are among the main hosts of entomophthoralean
fungi. In a seven year survey for the resources of this fungal group in
China, two new pathogen fungi were found. The first one was a patho-
gen of green bottle flies in northwestern China and could be attributed to
the genus Pandora. The second one caused an epizootic in a chironomid
midge population and was a member of the genus Erynia. They did not
agree with the descriptions of any published species of the respective
genera and are described here as two new species.
MATERIALS AND METHODS
The methods of Li and Humber (1984) were used for morphological
observation in glass slide material preparation. Lactophenol-cotton blue
(LPCB) and lactophenol-aceto-orcein (LPAO) (Keller and Wilding, 1985)
were used for staining.
RESULTS AND DISCUSSION
Some cadavers of green bottle flies, Lucilia sp., were found on leaves of
persimmon trees in Lintong, Southeastern Shaanxi Province, in north-
western China on 22 September 1991. The cadavers were attached to the
leaves by numerous rhizoids. The creamy hymenia emerged from inter-
segmental membranes. The microscopic appearance of the specimens
showed obvious characteristics of the genus Pandora Humber (1989): coni-
dia were uninucleate and bitunicate, cystidia were 2 times the thickness of
conidiophores, and the rhizoids had discoid holdfasts and were 2-3 times
the thickness of conidiophores. The species was not attributable to any
described species of this genus and is described here as new species:
PANDORA SHAANXIENSIS Fan & Li, sp. nov. [Figs. 1-5]
CONIDIA PRIMARIA uninucleata, bitunicata, incolorata, obovoidea,
papillis truncatalae, plerumque symmetrica, 30.3—43.2 xX 12.5-19.8 Um
(med. 32.743.0 x 15.0£1.2 um; L/D = 1.7-3.4). CONIDIA SECUN-
DARIA obovoidea sed minora, 13.2-16. 8 X 9.6—-12.0 Um (med. 14.9+0.9
x 10.1+0.8 um, L/D = 1.2-1.7). Capilliconidia desunt. CORPORA
HYPHALIA magna, ovoidea vel sphaerica, 79-5—-112.2 X 37.4-74.8 Um,
vel clavata, 65.4-107.5 x 14.0-23.4 um. CONIDIOPHORA bifurcata,
6.9-9.2 um in diam. CYSTIDIA aseptata, robusta, 13.8-16.1 Um in diam.
RHIZOIDEA hyphoidea, desinentia in haptera discoidea, 16.7—31.7 Um
in diam. SPORAE PERDURANTES desunt.
309
FIGURES 1-5. Pandora shaanxiensis. 1. Primary and secondary coni-
dia, X260. 2. Conidiophores, X440. 3. Hyphal bodies and conidiophores
arising from the hyphal bodies, X250. 4. Rhizoids with disc-like holdfast,
x210. 5. Conidiophores and a cystidium, X440.
310
PRIMARY CONIDIA (Fig. 1) uninucleate, bitunicate, colorless, ob-
ovoid, with nearly rounded apex and a slightly truncate papilla, mostly sym-
metnc, '30/3-43.2)'% 125-19 8) tn (averaging) 32, 743.0) x) 15\0g1) Zelim
L/D = 1.7-3.4, averaging 2.4+0.3; n = 100), with cytoplasm containing
several vacuoles, forcibly discharged by papillar eversion. SECONDARY
CONIDIA (Fig. 1) similar to primary conidia but smaller; no capilliconidia
formed. HYPHAL BODIES large, ovoid to subspherical, 79.5-112.2 x
37.4-74.8 [tm, or clavate, 65.4-107.5 xX 14.0-23.4 um. Conidiophores
arise from these hyphal bodies; cytoplasmic transfer to the conidiophores
leaves empty saccular cells (Fig. 3). NUCLEI elliptical to subglobose,
centrally located, 7.0-12.2 X 4.5-8.4 Um in primary conidia. CONIDIO-
PHORES bifurcately branched (Figs. 2,5), determinate, 6.9—9.2 Um wide,
covering intersegmental membranes. CYSTIDIA (Fig. 5) aseptate, rarely
branched, robust, 13.8-16.1 Um wide. RHIZOIDS (Fig. 4) hypha-like,
terminating in discoid holdfast, usually in bundle of several rhizoidal
threads, 16.7—31.7 Um thick. RESTING SPORES not observed.
Holotype: ACAFP 11027; adult of green bottle fly; deposited at herbari-
um of Forest Protection, Department of Forestry, Anhui Agricultural
University. Collected by Dahong Yang.
Type Host: Lucilia sp. (Diptera: Calliphoridae)
Type Locality: Lintong, Shaanxi; 22 September 1991.
There are no species in the genus Pandora which have conidial mea-
surements close to those of P. shaanxiensis. Very few specimens were
collected in and around the type locality, thus suggesting that the host flies
were infected during an enzootic.
The second species caused an epizootic in a chironomid midge popula-
tion. Greyish green hymenia appeared first from intersegmental mem-
branes of the midge adbomens, and eventually covered almost the whole
bodies of the hosts.
PANDORA CHIRONOMIS Fan & Li, sp. nov. [Figs. 6-9]
CONIDIA PRIMARIA uninucleata, bitunicata, incolorata, anguste ob-
ovoidea, plerumque asymmetrica interdum parum curva, 29.2-60.6 X
10.4—21.9 lum (med. 41.14 5.7 — 48.0410.1 X 13.541.9 — 17.143.8 Um;
L/D = 2.1-4.3, med. 2.5+0.3-3.6+ 0.5), papillis angustis, longis vel
parum rotundatis, collariatis.s CONIDIA SECUNDARIA similia conidi-
orum primariorum sed angustora, 28.2—37.9 x 8.0-10.4 Um (med. 33.44
FIGURES 6-9. Erynia chironomis. 6. A midge cadaver with thick
hymenium on the abdomen, X7. 7. Rhizoids and conidiophores, X80. 8.
Primary conidia, X360. 9. Cystidia and conidiophores, X130.
ez
3.4 X49 .920 9 my LAD =) 2.93.9) Svelisphaenca: pyntonniaet 4-20. 9a
b2-1553/ im (nied 118.7410 xX 13. 8L0s80 Ning L/D =e 14) capil
conidia et conidia coronata desunt. CORPORA HYPHALIA clavata, 6.3-
10.4 Um in diam. CONIDIOPHORA ramosa vel simplicia, 8.4-13.6 Um
in diam. CYSTIDIA abunda, longa, robusta, aseptata, 21.7-38.4 [um in
diam., basaliter, usque ad 668 Um longa. RHIZOIDEA abunda, mono-
hyphoidea, 14.5-24.3 Um in diam. SPORAE PERDURANTES desunt.
PRIMARY CONIDIA (Fig. 8) uninucleate, bitunicate, colorless, narrowly
obovoid, usually asymmetric and sometimes curved, with largest diameter
in the apical half, apex broadly rounded or somewhat truncate, tapering to a
narrow, long, triangular or slightly rounded and collared papilla; 29.2-60.6 x
10.4-21.9 um (averaging 41.4+5.7 — 48.0410.1 « 13.5+1.9 — 17.143.8
Lim; LAD tie! 2514.3) averaging 22520). 08 8:00.51 a) ace O) ae
cytoplasm containing one to several large vacuoles; forcibly discharged by
papillar /eversion, SECONDARY CONIDIA (Figs 39) 1 O)esimnitarero
primary conidia but narrower, 28.2—37.9 X 8.0—10.4 (averaging 33.4+3.4
* "9 9t0'9 um; L/D: = 2:7-3.9, averaging 3420 4). or sphericalgte
pyniform, 17.4—20.5 xX 12.2-15.3 um (averaging 18.7+1.0 xX 13.8+0.8 Lm;
L/D = 1.2-1.4, averaging 1.40.1); no capilliconidia or tetraradiate
conidia formed. HYPHAL BODIES clavate, 6.3-10.4 lm wide. NU-
CLEI elliptical, 6.3—20.9 x 3.5-7.0 Um in primary conidia. CONIDIO-
PHORES branched or simple, 8.4-13.6 Um wide, usually forming
continuous and thick greyish-green hymenia eventually covering the whole
bodies of the hosts. CYSTIDIA (Fig. 9) abundant, long, robust, aseptate,
unbranched, 14.6—22.8 Um wide in the middle and 21.7—38.4 Um wide at
the level of the conidiophores, up to 688 Um long. RHIZOIDS (Fig. 7)
abundant, robust, long, monohyphal, 14.5—24.3 Um wide without differ-
entiated holdfast. RESTING SPORES not observed.
Holotype: ACAFP 10523; adult of midge; deposited at herbarium of
Forest Protection, Department of Forestry, Anhui Agricultural Univer-
sity. Collected by Zengzhi Li.
Type Host: Chironomus sp. (Diptera: Chironomidae)
Type Locality: Hefei, Anhui; 23 May 1991.
This pathogen caused very high mortality (>80%) of the midges in a
little pool during a rainy season which lasted for almost a continuous month.
The macroscopic and microscopic appearance resemble those of Erynia
conica (Nowakowski) Remaudiére & Hennebert and E. rhizospora (Thax-
ter) Remaudiére & Hennebert. E. chironomis, however, has the largest
Sit
diameter of conidia in the apical half and is usually much narrower although
its length is within that of E. conica (Thaxter, 1888; Gustafsson, 1965;
Keller, 1991). The new species can be distinguished from E. rhizospora
(Thaxter, 1888; Keller, 1991) by its much longer and narrower conidia and
by its shorter papillae. The conidia of E. chironomis are also similar to those
of E. aquatica (Anderson & Anagnostakis) Humber, but those of the latter
are much shorter and narrower and usually have only a single large vacuole
(Anderson and Ringo, 1969).
The generic name Erynia has been confusing. Nowakowski (1881)
published it invalidly and, in fact, rejected it (Nowakowski, 1883). Batko
(1966) validated this name at the subgeneric level 83 years later, but
Remaudiére and Hennebert (1980) used the validated Erynia at the same
(generic) rank with Zoophthora. Humber and Ben-Ze’ev (1981), however,
emended Erynia Nowakowski to cover all species of Zoophthora Batko.
Ben-Ze’ev and Kenneth (1982) used this generic name in the same,
expanded sense. Humber (1989) proposed a new classification in which the
emended Erynia was divided into 4 genera — Erynia, Furia, Pandora, and
Zoophthora. Keller (1991) accepted the revision of Remaudiére and
Hennebert (1980) and rejected both the emendation by Humber and Ben-
Ze’ev (1981) and the separation by Humber of the four genera. Based on
the taxonomic work leading to the descriptions of Furia shandongensis
(Wang et al., 1994) and of Erynia chironomis and Pandora shaanxiensis here,
the revision by Humber (1989) is found to be reasonable, practical, and
acceptable.
ACKNOWLEDGMENTS
We are particularly grateful to Dr. R. A. Humber for his critical review
of the manuscript, and for its final preparation and submission.
LITERATURE -CLLUED
Anderson, J. F. and S. L. Ringo. 1969. Entomophthora aquatica sp. n.
infecting larvae and pupae of floodwater mosquitoes. J. Invertebr. Pathol.
13: 386-393.
Batko, A. 1966. On the subgenera of the fungus genus Zoophthora Batko
1964. Acta Mycol. 2: 17-21.
Ben-Ze’ev, I. and R. G. Kenneth. 1982. Features-criteria of taxonomic
value in the Entomophthorales. II. A revision of the genus Erynia
Nowakowski 1881 (= Zoophthora Batko, 1964). Mycotaxon 14: 456-
475.
314
Gustafsson, M. 1965. On species of the genus Entomophthora in
Sweden. I. Classification Lantbrukshogsk. Ann. 31 103-212.
Humber, R. A. 1989. Synopsis of a revised classification for the Ento-
mophthorales (Zygomycotina). Mycotaxon 34: 441-460.
Humber, R. A. and I. Ben-Ze’ev. 1981. Erynia (Zygomycetes: Ento-
mophthorales) emendations, synonymy and transfers. Mycotaxon 13:
506-516.
Keller, S. 1991. Arthropod-pathogenic Entomophthorales of Switzerland.
II. Erynia, Eryniopsis, Neozygites, Zoophthora, and Tarichium. Sydowia
43: 39-122.
Keller, S., and N. Wilding. 1985. Entomophthora brevinucleata sp. nov.
(Zygomycetes, Entomophthoraceae), a pathogen of gall midges (Dip.:
Cecidomyiidae). Entomophaga 30: 55-63.
Li, Z. and R. A. Humber. 1984. Erynia pieris (Zygomycetes: Ento-
mophthoraceae), a new pathogen of Pieris rapae (Lepidoptera: Pieri-
dae): description, host range, and notes of Erynia virescens. Can. J. Bot.
62: 653-663.
Nowakowski, L. 1881. O grupie owadomorkow (Empusaceae). Dzienn.
HI Zjadu Lek. Pyzyr. Polsk. Krakow, Sekt. Bot. 6: 67-68.
Nowakowski, L. 1883. Entomophthoreae. Pyzycznek do znajamosci
pasorzytnych grzybk6ow sprawiajgcych pomor owadow. Pamietn. Wydz.
Akad. Umiej. w Krakow 8: 153-183.
Remaudiére, G. and G. Hennebert. 1980. Révision systématique de
Entomophthora aphidis Hoffm. in Fres. Description de deux nouveaux
pathogénes d’aphides. Mycotaxon 11: 323-338.
Thaxter, R. 1888. The Entomophthoreae of the United States. Mem.
Boston Soc. Nat. Hist. 4: 133-201.
Wang, W., W. Lu, and Z. Li. 1994. Furia shandongensis, a new patho-
gen of earwigs. Mycotaxon 50: 301-306.
MYCOTAXON
Volume L, pp. 315-321 January-March 1994
A NEW CORTINARIUS FROM A MATURE ASPEN STAND
IN MONTANA
CATHY CRIPPS AND ORSON K. MILLER, JR.
Department of Biology
Virginia Polytechnic Institute and State
University, Blacksburg, VA 24061
ABSTRACT
Cortinarius hedyaromaticus, a large, white Phlegmacium
with a strong, sweet odor, is described as a new species from
a mature stand of Populus tremuloides in southwestern Montana.
Key Words: aspen, Cortinarius, Phlegmacium, Populus
tremuloides, Montana.
INTRODUCTION
Large clonal stands of aspen (Populus tremuloides Michx.)
are a prominent feature of the landscape in western North
America, especially in montane regions associated with the
Rocky Mountains. Although aspen is usually considered a
pioneering species in this area, eventually being replaced by
conifers, it may persist to form mature stands. An unknown
large, whitish, sweet-smelling Cortinarius with a greasy cap
was discovered in one such stand during a survey to document
the ectomycorrhizal fungi associated with aspen in
southwestern Montana (Cripps 1992). The stand is on the floor
of a glaciated valley at an elevation of 1900 m. in Park
County, Montana, 5 miles north of Yellowstone National Park.
Although the stand is nearly pure aspen, engelmann spruce
(Picea engelmanii Parry ex. Engelm.), douglas-fir (Pseudotsuga
menziesii Mirb.Franc), and lodgepole pine (Pinus contorta
Dougl. ex Loud) have invaded in some areas, and willow and
alder are found in the wetter areas. These are all
potentially ectomycorrhizal trees, but aspen is the most
likely host. Ten of the largest trees on the site were cored
316
and ages ranged from 56 to 114 years, averaging 81 years.
This is a productive site with a deep, gravelly loam soil of
glacial till, and has remained undisturbed by fire for many
years.
Sporocarps of the Cortinarius were found in the same
local area in late summer in 1991, and 1992 on this site, and
representative specimens were mailed to Dr. Meinhard Moser who
agreed this was a new taxon.
Methods: Collections were photographed on the site, smell and
taste recorded, and the sporocarps wrapped in wax paper.
Potential mycorrhizal hosts were noted. Morphological
descriptions were made on fresh material. Specimens were
dried for microscopic examination at a later date. All
sections were prepared in 3% KOH unless otherwise noted.
Collections are in the Massey Herbarium (VPI), Blacksburg,
Virginia:
TAXONOMY
Cortinarius hedyaromaticus sp. nov. C. Cripps and O.K. Miller
Livustrat rons: higge. L=7..
Pileus: 7-15 cm latus, hemisphaericalis vel late convexus,
rasilis, in juventute sebosus, aridus, saepe areolatus in
aetate; margo persistenter involutus, interdum appendiculatus,
bubalinus sed in aetate corylinus, si contusus porphyreus,
rare violaceo-griseo tinctus. Lamellae subdecurrentes,
confertae, parum latae, in juventute albidae, deinde
brunnneae; margines brunneo-vinosi sSi_ contusi. Stipes
variabilis, 7-11 cm longus, -2.5 cm latus, aequalis vel ad
basem auctus deinde abrupte decrescens usque ad punctum
vinosum si contusum, bubalinus aridus fibrillis sparsis; apex
albus pulveraceus, in aetate interdum lilacinus super zonam
annularem, caro lenta. Velum partiale cortinatum vel paene
membranaceum, album, superiorem zonem annularem relinquens.
Contextus firmus, albus, interdum ferrugineus si contusus.
Odor) dulcis, saepe fortis. Sapor mitis vel parum amarus.
Sporae 9.5-11(-13) x 5.5-6.5(-7.5) wm diametro, verrucosae
Ssubamagdaliformes, extremo obtuso. Depositum sporarum
brunneum. Pileipellis mixocutis est hypharum filamentosarum.
Trama lamellaris parallela cellularum 7-15 wm latarum.
Cystidia /absentia-#y Typum legit: CLO 316¢q,.28 August yet oie
Cinnabar Basin, Park Co., Montana.
Pileus: up to 15 cm broad, at first hemispherical with
heavily inrolled margin becoming broadly hemispherical (Figs.
3 & 4), convex or domed with wavy or inrolled margin which is
sometimes areolate or rimose, greasy at first, becoming smooth
and dry, at center often coarsely areolate in age or on
drying, cream buff (whitish)/» or< pinkish=buis “atiptiesy,
bruising rusty brown especially at the margin (reminiscent of
a Hebeloma), occasional appendiculate pieces of the white
partial veil (Fig. 6), maturing to light brown and remaining
so when dried, localized areas often with steel grey, blue
317
grey, grey brown, or grey violet tints at different maturation
stages. Lamellae subdecurrent, crowded, medium broad, pale
cream-buff to clay color young becoming light cocoa then
brown, edges bruising vinaceous-brown, violet-gray to steel-
gray. Stipe extremely variable, 7-11 cm long, up to 2.5 cm
broad, equal, enlarging somewhat toward the base and then
abruptly tapered and pointed at the base (which may bruise
vinaceous), ary, whitish cream at first with sparse rusty
fibrils and a few very pale yellow patches at apex, apex often
powdery, with steel-gray to gray-violet tints above superior
annular zone in older specimens, on bruising, or on drying
(Fig. 5), stipe flesh extremely firm. Partial veil cortinous
to nearly membranous (Fig. 7), white, soon gone leaving
Superior rusty zone. Context firm, white, brown around worm
holes, staining buff to vinaceous or red-brown at stipe apex,
pale yellow in ammonia, negative in FeS04. Taste mild to
slightly bitter. Smell strong, sweet like vanilla cake batter
(rarely faint), then sickenly sweet on drying!
Pileipellis: a mixocutis, upper layer of gelatinized, hyaline,
filamentous hyphae, 3.5-4.2 um wide, below a golden layer of
hyphae, 3.5-5.0(-7) um wide, some with pigmented walls, some
hyaline, some with incrustations, clamps present. Pileus
trama of broadly filamentous, inflated hyphae, 7.6-13 (-20)
um wide, clamps scattered. Lamellar trama parallel, of thin-
walled hyphae 7-15 um wide. Subhymenium a narrow layer of
hyphae 5-8 um in diameter. Basidia, 4-spored, clavate, 30-40
Shel OU, volLanps at. base.) (Fig.: .Wh)i..) Partial.~ veil,..of
filamentous hyphae, 3-5 um wide, clamps frequent.
Basiqiospores:: 975-11 (-13) x 5.5=-605)(=7..5)n ume eD™.=) 10. 4 .x).6,,0
um, average Q = 1.7, verrucose, subamygdaliform to almost
almond-shaped, with slightly flattened, blunt obtuse end (Fig
2). Spores brown in mass, not rusty-brown.
Habitat & distribution: terrestrial, often in clumps or ina
ring, beneath mature aspen (Populus tremuloides) nis 9)
southwestern Montana. Fruiting in late summer.
Material examined: United States. Montana, Park Co, Cinnabar
basin: 28VAugust, (1991, 1CLC1 316. (holotype): 11. August, .1991,
CLE E260 lS JULY ,, 11992.
Observations: The greasy cap and dry stipe place this
Cortinarius firmly in the subgenus Phlegmacium. The stirps
Rapaceum in Section Phlegmacium (Moser 1960) which is
characterized by a white, whitish, to cream-ochraceous cap
color, as well as clay-colored lamellae when young, best fits
our, taxon.) Within: this group; Cc. lustratus Fr.,.has a similar
color and looks related, but is small, has smaller spores
(6,.5->80e) 3..5-4..5 pm), .andsa tfloury,.smeli. (iC. aleuriosmus. R.
Mre has similar colors but a rather strong farinaceous odor,
and a marginately bulbous base. The, (same: sis. true (TOL. £..
osmophorus P.D. Orton, which is described with bluish shades
in the stipe, a taxon Moser (pers. written comm.) believes to
be a synonym of C. aleuriosmus, where slight bluish shades can
318
occur. C. amarescens (Mos.) Mos. is similar but has an abrupt
bulb, strongly bitter “flesh (not in. ours) j and is a veces
robust species with a cap 3-5.5 cm in diameter, stipe 6-7.5
cm long, 9-15 mm wide with bulb 15-20 mm wide. Our stipe is
7-1 tvem long, sup Conic. > Tem DEoaas Flora Agaricina Danica
(Lange 1938) plate 82. fig. B (C. talus ss Lange), is in the
related group with similar coloration, a viscid cap 4-7 cn,
Similar spores 8.5-10.2(-11) x 5-5.5 um, but the bulb is
marginate, abrupt and rounded unlike ours, and the taste is
bitterish. Moser considers C. talus ss Lange above as a
synonoym of Phlegqmacium amarescens (Mos.) Mos. (Moser 1960,
p. 146). Lange (1938)) says the ‘taxon’ is “very close tome:
multiformis but paler, more viscid and somewhat bitter." C.
multiformis (Fr.) Fr’. (differs from our’ taxon in an?) ochre—
pallid to yellow-brown cap color and the presence of a bulbous
base, but has similar spores and a similar sweet honeylike
odor (illustrations: Dahnke and Dahnke 1979, Bresadola 1982,
Ricken 1915).
Our taxon would be placed in subsection Multiformes if
odor were the only consideration, but is placed in Rapacei for
other characters (Moser 1978). Also in subsection
Multiformes, Phlegmacium talus (Fr.) Blytt (C. talus Fr.) as
described in Die Gattung Phlegmacium (Moser 1960) is
characterized by an ochre-brown to reddish-brown cap similar
to the more highly colored caps in this subsection. It is a
large species with a slightly inrolled margin (ours is
distinctly inrolled), the smell is sweet (orange blossom to
Kunsthonigartig--honeylike) similar to ours, spores 8.5-10(-
11) x 4.8-5.5 wm are somewhat small in size, overlapping ours,
but ait has an, abrupt bulbvuniike ours. ~P- talus (Fr. jee
occurs in beechwood on calcareous soil in Europe which is
different from ours. €. “ochropallidus ~Rv/°Hry.” “an serne
Multiformes group has a pallid cap similar to our taxon, but
is less robust, cap 4=7 cm, has a bluntly abrupt bulb,;"/ana
Spores 11-11.5 x 5.5-6 um somewhat larger than ours, with no
odor.
In summary, Moser in Singer (1986) places these related
taxa in Phlegmacium Subsection Multiformes Henry ex Moser.
Stirps Multiformis contains C. multiformes (Fr. ex Secr.) and
Cc. talus Fr., and Stirps Rapaceus contains C. amarescens (Mos.
ex Mos.) Moser, C. rapaceus Fr., and C. albidus Peck. The
latter stirps best fits our taxon because Vof™ cap color,
however, other taxa with a sweet odor are found in Stirps
Multiformis.
There’ is "Littte doubt ‘that’ our taxon’ ts part or wunus
complex of species. It is the only species in the complex
associated with Populus tremuloides. The closest’ taxon an
size, coloratvon, and Spore’sizevis Cc: ‘talus (rr) Stytu sand
especially noted is the sweet smell. cC. hedyaromaticus has
a very ‘pallid; (almost “‘white), colored) "cap;*"ar distinct,
inrolled margin, is the most robust species in the complex,
has large spores, a sweet smell, mild taste (only slightly
bitter at times), and is the only taxon of the group reported
to be associated with Populus tremuloides in North America.
319
Figs. 1-3. Cortinarius hedyaromaticus. 1. basidia 30-40 x 8-
10 pm. 2. basidiospores 9.5-11(-13) x 5, OO sO 75 Oo) um.
Scale bar = 10 pm. 3. sporocarp 1/4 natural size.
320
Figs. 4-7. Cortinarius hedyaromaticus. 4. inrolled margin and
enlarged non-marginate stipe base (3/4 actual $126). 5. older
specimens showing gray-violet statining reaction at stipe apex
(1/2 natural size). 6. appendiculate margin and subdecurrent
gills (1/2 .nacural size). 7. membranous veil on younger
specimen (1/2 natural size.
ai
ACKNOWLEDGEMENTS
We would like to thank Dr. M. Moser for his assistance and
for reviewing this article, Dr. T. Macadoo for the Latin
diagnosis, Dr. R. Treu for taxonomic consultation, D. Bachman
for assistance in field work, as well as The Virginia Museum
of Natural History for logistical support.
LITERATURE CITED
Cripps, C. L. 1992. Aspen mycorrhizae: ecology, syntheses, and
growth studies. M.S. Thesis, Virginia Polytechnic Inst.
and State Univ., Blacksburg), Va.)\ 157 p.
Bresadola, J. 1927-1933, 1941 (New Edition 1982). Iconographia
Mycologica. Vol IV. Mediolani.
Dahnke, R. and S. D&ahnke. 1979. 700 Pilze in Farbfotos. AT
Veriag Aarau, Stuttgart...671 p.
Lange, J. E. 1938. Flora Agaricina Danica III, Kopenhagen,
Denmark. 96 p.
Moser, M. 1960. Die Gattung Phlegmacium. Verlag Julius
Klinkhardt, Bad Heilbrunn (Obb.), Regensburg, Germany.
440 p.
Moser, M. 1978. Keys to Agarics and Boleti. Gustav Fischer
Verlag, stuttgart, Germany. 535 p.
Ricken, A. 1915. Die Blatterpilze (Agaricaceae). Theodor
Oswald Weigel, Leipzig. 480 p.
Singer, R. 1986. The Agaricales in modern taxonomy. Koeltz
Scientific Books, Koenigstein, Germany. 981 p.
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MY COTAXON
Volume L, pp. 323-330 January-March 1994
THE LICHEN FLORA OF ROCK CANYON, UTAH COUNTY,
UTAH
Ted O. Ririe!, Larry L. St. Clair!, and Clayton C. Newberry
1 Department of Botany and Range Science, Brigham Young
University, Provo, UT 84602 2 Department of Integrative Biology,
University of California, Berkley, CA 94720
ABSTRACT
Sixty-three lichen species in thirty genera are reported for
Rock Canyon, a steep, west-facing canyon in the Wasatch
Mountains of central Utah. Dimelaena thysanota, Acarospora
boulderensis, Physcia dimidiata and Rinodina turfacea are
reported as new to Utah.
INTRODUCTION
Literature review of Utah lichenology
Utah lichenology began with Edward Tuckerman's 1871 report
of 27 species and 8 varieties from various locations in the state
(Tuckerman 1871). Until recent times, however, botanists and
other collectors paid little attention to the lichen flora of Utah.
Possibly the relative scarcity of showy macrolichens in the state
discouraged serious floristic investigation.
Over eighty years after Tuckerman's initial report, Seville
Flowers (1954) published a list of 127 lichen species in 27 genera.
Flowers was a bryologist and his lichen collections were merely
incidental to his statewide bryological investigations; but his
collections indicated that Utah had a rich microlichen flora in need
of more serious study.
Harry Nielsen (1960) completed "Lichens of the Wasatch
Mountains in Central Utah" for his masters thesis at Brigham
Young University. Nielsen's thesis included the first keys and
formal descriptions of Utah lichens, and included 52 species and 2
varieties in 17 genera, with 33 species reported as new for Utah.
Anderson and Rushforth (1976) published the first of many
studies on cryptogamic soil crusts in Utah's arid to semiarid
habitats. They reported 17, mostly terricolous lichen species, in
324
11 genera. As part of an air quality survey commissioned by the
National Park Service for the southern portion of Zion National
Park, Nash and Sigal (1981) reported 159 lichen species in 53
genera, 4 of which were new to Utah.
St. Clair (1984) studied lichen communities along an alpine
tundra ridge in the Uinta Mountains of northeast Utah, and
reported 66 species, 16 of which were new to the state. St. Clair
and Warrick (1987) reported Acarospora nodulosa (Duf.) Hue. var.
nodulosa from gypsiferous soils in southern Utah as a new record
for North America.
Recently, a comprehensive literature review of the lichens of
Utah, based on reports in 44 papers, was published (St. Clair et al.
1991). A total of 85 genera, 318 species, and 11 varieties have
been reported for Utah. Based on lichenological investigations
throughout the state and in conjunction with the publication of the
catalog, Newberry and St. Clair (1991) reported an additional 72
species in 45 genera as new to the state. This brought the total
number of Utah lichens to 390.
Newberry (1991) described the lichen flora of the Uinta
Moutains, and reported 291 taxa in ninety-five genera from his
study. St. Clair and others are currently completing a floristic
survey of the lichens of Timpanogos Cave National Monument,
Utah and to date have identified 72 species in 29 genera for the
monument.
Few lichen ecology studies have been conducted in the state of
Utah. One of the first papers on this subject dealt with lichen
community structure in Zion National Park (Rushforth et al 1982).
Cluster analyses and Shannon-Weaver diversity indices were used
to analyze field data and characterize lichen community structure
in the park. St. Clair's thesis, referenced above, characterized
lichen communities along an elevational gradient in the Uinta
Mountains (St. Clair 1984). This study assessed the influence of
vascular plant cover, rock cover, soil development, grazing and
snow-melt patterns on the distribution of alpine tundra lichens. A
study considering the influence of microhabitat on species
diversity, distribution and abundance of corticolous lichens in Zion
National Park was completed in the mid 1980s (St. Clair et al
1986).
Description of study site
Rock Canyon is located in Utah County, 63 km south of Salt
Lake City, UT. Along with numerous side canyons, Rock Canyon
cuts 4 km through the Wasatch Mountains from east to west and
O20
opens to the west at approximately 40° 16' north latitude, 111°
37' west longitude. Elevation in the canyon bottom ranges from
2134 m in the upper gorge to 1524 m at the mouth of the canyon.
Sequentially from the canyon's river bed the stratigraphy
consists of Tintic Quartzite, Pre-Cambrian Tillite, and Great Blue
Limestone.
Vascular plant communities in Rock Canyon vary with
elevation and proximity to water. The Mountain brush
community in the lower elevations includes gambel oak (Quercus
gambelii), bigtooth maple (Acer grandidentatum), serviceberry
(Amelanchier alnifolia), two species of mountain mahogany
(Cercocarpus ledifolius and Cercocarpus intricatus), and big
sagebrush (Artemisia tridentata). The riparian community
includes narrowleaf cottonwood (Populus angustifolia), water
birch (Betula occidentalis), thinleaf alder (Alnus incana), box elder
maple (Acer negundo), red-osier dogwood (Cornus sericea),
horsetail (Equisetum hyemale), willow (Salix scouleriana), and
some big sagebrush (Artemisia tridentata). The upslope juniper
community includes Utah juniper (Juniperus osteosperma), and
big sagebrush (Artemisia tridentata), usually on steep, well-
drained exposed slopes. Middle elevations are characterized by
quaking aspen (Populus tremuloides), big sagebrush (Artemisia
tridentata), and lodgepole pine (Pinus contorta). Higher elevations
are characterized by englemann spruce (Picea engelmannii),
subalpine fir (Abies lasiocarpa) and lodgepole pine (Pinus
contorta).
Uinta National Forest administers Rock Canyon, protects it as a
critical watershed, prohibits motorized vehicle use in the canyon
and maintains a primitive foot trail. In recent years civic groups
have proposed that Rock Canyon be managed more
comprehensively for natural research and low-impact recreation.
METHODS
As lichen distribution is influenced by substrate, moisture and
sunlight, all available substrates and habitats in the canyon have
been carefully sampled. Small amounts of each species from each
substrate and habitat were collected. Specimens were removed
directly from the substrate where possible, or depending on the
species, sometimes attached to small pieces of bark, soil, or rock.
Specimens were placed in carefully labeled paper sacks and
taken to the BYU Herbarium of Nonvascular Cryptogams, where
they were curated and placed in permanent herbarium packets
326
labeled with site, habitat and substrate information. Species were
identified using standard lichen keys and taxonomic treatises.
Standard chemical spot tests and, where necessary, thin-layer
chromatography techniques were used to finalize species
identifications.
RESULTS
Sixty-three lichen species in thirty genera are reported from
Rock Canyon. Acarospora boulderensis, Dimelaena thysanota,
Physcia dimidiata, and Rinodina turfacea are reported as new for
Utah. Specimens are deposited in the Herbarium of Nonvascular
Cryptogams at Brigham Young University (BRY).
Following is an alphabetic list of all lichen species
collected in Rock Canyon. The species are followed by the BRY
herbarium number, substrates, and relative abundance. Species
marked with an asterisk are new state records.
*ACAROSPORA BOULDERENSIS Magnusson. BRY C21815. Rare. On
quartzite.
ACAROSPORA GLAUCOCARPA (Ach.) Ko6rber. BRY C21883. Rare. On
limestone.
ACAROSPORA STRIGATA (Nyl.) Jatta. BRY C21884. Rare. On limestone.
ASPICILIA CALCAREA (L.) Mudd. BRY C21885, BRY C21886, BRY
C21887. Rare to common. On limestone.
ASPICILIA CINEREA (L.) Korber. BRY C22391. Rare to common. On
quartzite.
ASPICILIA DESERTORUM (Krempelh.) Mereschk. BRY C21888, BRY
C21889. Rare to common. On limestone.
CALOPLACA CITRINA (Hoffm.) Th. Fr. BRY C21890. Rare. On
limestone.
CALOPLACA HOLOCARPA (Hoffm.) Wade. BRY C21891. Common. On
bark of narrowleaf cottonwood.
CALOPLACA TRACHYPHYLLA (Tuck.) Zahlbr. BRY C21892. Common. On
limestone.
CANDELARIELLA DEFLEXA (Nyl.) Zahlbr. BRY C21893, BRY C21894.
Common. On bark of narrowleaf cottonwood, and gambel oak.
CANDELARIELLA ROSULANS (Miill. Arg.) Zahlbr. BRY C21895. Common.
On quartzite.
CATAPYRENIUM GRANULOSUM (B. de Lesd.) Thomson. BRY C21896.
Rare. On limestone.
CATAPYRENIUM LACHNEUM (Ach.) R. Sant. BRY C21897. Rare. On soil
over limestone.
oan
COLLEMA COCCOPHORUM Tuck. BRY C21898. Rare. On limestone.
COLLEMA FLACCIDUM (Ach.) Ach. BRY C21899, BRY C21900. Rare to
common. On limestone.
COLLEMA POLYCARPON Hoffm. BRY C21901. Rare. On limestone.
DERMATOCARPON INTESTINIFORME (Korber) Hasse. BRY C21902, BRY
C22390. Rare. On quartzite.
DERMATOCARPON MINIATUM (L.) Mann. BRY C21903, BRY C21904.
Common to abundant. On limestone.
DERMATOCARPON RETICULATUM Magnusson. BRY C21905. Common. On
limestone.
DIMELAENA OREINA (Ach.) Norman. BRY C21906. Common. On
quartzite.
*DIMELAENA THYSANOTA (Tuck.) Hale & Culb. BRY C21677. Rare. On
quartzite.
DIPLOTOMMA ALBOATRUM (Hoffm.) Flotow. BRY C21907. Rare. On
limestone.
FULGENSIA FULGENS (Swartz) Elenkin. BRY C21908. Rare. On soil
over limestone.
GONOHYMENIA NIGRITELLA (Lettau) Henssen. BRY C21909. Rare. On
limestone.
HYPERPHYSCIA ADGLUTINATA (Flérke) H. Mayrh. & Poelt. BRY C21940.
Common. On bark of gambel oak.
LECANORA DISPERSA (Pers.) Sommerf. BRY C21910. Rare. On
quartzite.
LECANORA HAGENI (Ach.) Ach. BRY C21911, BRY C21912. Common.
On bark of narrowleaf cottonwood, and gambel oak.
LECANORA MURALIS (Schreber) Rabenh. BRY C21913. Rare on
lignum. This is an unusual substrate for L. muralis. Ordinarily
this species occurs on rock but in western North America it
occasionally appears on lignum of various conifers.
LECANORA THALLOPHILA Magnusson. BRY C21914. Rare. On
Dermatocarpon intestiniforme.
LECIDEA LEUCOTHALLINA Arnold. BRY C21915. Common. On
quartzite.
LECIDEA TURGIDULA Fr. BRY C21916. Rare. On bark of narrowleaf
cottonwood.
LECIDELLA STIGMATEA (Ach.) Hertel & Leuck. BRY C21939. Rare to
common. On limestone.
LEPRARIA INCANA (L.) Ach. BRY C21917. Rare. On moss.
MEGASPORA VERRUCOSA (Ach.) Hafellner & Wirth. BRY C21918, BRY
C21919. Rare to common. On soil, and moss over quartzite.
PELTIGERA CANINA (L.) Willd. BRY C21920. Common. On moss over
rock.
328
PHAEOPHYSCIA CILIATA (Hoffm.) Moberg. BRY C21921. Rare. On bark
of douglas fir.
PHAEOPHYSCIA ENDOCOCCINA (Korber) Moberg. BRY C21922. Rare. On
quartzite.
PHAEOPHYSCIA NIGRICANS (Floérke) Moberg. BRY C21923. Rare to
common. On bark of narrowleaf cottonwood.
PHAEOPHYSCIA ORBICULARIS (Necker) Moberg. BRY C21924, BRY
C21925, BRY C21926, BRY C21927, BRY C21928, BRY C21929,
BRY C21930, BRY C21931, BRY C21932. Common to abundant.
On bark of narrowleaf cottonwood, bigtooth maple, shaded
limestone, limestone, quartzite, moss over quartzite, moss over
limestone, moss.
PHYSCIA CAESIA (Hoffm.) Fiimmr. BRY C21933, BRY C21934. Rare to
common. On moss over quartzite, quartzite.
*PHYSCIA DIMIDIATA (Arnold) Nyl. BRY C21935. Rare. On bark of
gambel oak.
PHYSCIA DUBIA (Hoffm.) Lettau. BRY C21936, BRY C21937, BRY
C21938. Common. On bark of bigtooth maple, quartzite, moss.
PHYSCIA TENELLA (Scop.) DC. in Lam. & DC. BRY C21939. Rare. On
limestone.
PHYSCONIA DETERSA (Nyl.) Poelt. BRY C21941. Rare. On moss over
quartzite.
PHYSCONIA GRISEA (Lam.) Poelt. BRY C21942, BRY C21943, BRY
C21944. Rare. On bark of gambel oak, moss.
PHYSCONIA MUSCIGENA (Ach.) Poelt. BRY C21946, BRY C21947, BRY
C21948, BRY C21949. Rare to common. On limestone, soil,
moss, moss over quartzite.
PSORA TUCKERMANII R. Anderson ex Timdal. BRY C21950. Common.
On soil over limestone.
RHIZOCARPON GEMINATUM Korber. BRY C21951, BRY C21952.
Common. On quartzite.
RHIZOPLACA MELANOPHTHALMA (DC. in Lam. & DC.) Leuck. & Poelt. BRY
C21953, BRY C21954. Rare to common. On limestone and
quartzite.
RHIZOPLACA PELTATA (Ramond) Leuck. & Poelt. BRY C21955. Rare to
common. On quartzite.
RINODINA BISCHOFFII (Hepp.) Massal. BRY C21956. Rare. On
limestone.
RINODINA PYRINA (Ach.) Arnold. BRY C21957, BRY C21958, BRY
C21959. Rare to common. On bark of bigtooth maple,
narrowleaf cottonwood and gambel oak.
*RINODINA TURFACEA (Wahlenb.) Korber. BRY C22392. Rare. On
bark of douglas fir.
o29
TONINIA CAERULEONIGRICANS (Lightf.) Th. Fr. BRY C21960, BRY
C21961. Rare. On soil, and moss over limestone.
UMBILICARIA PHAEA Tuck. BRY C21962. Common to abundant. On
quartzite.
UMBILICARIA TORREFACTA (Lightf.) Schrader. BRY C21963. Rare. On
limestone.
UMBILICARIA VIRGINIS Schaerer. BRY C21964, BRY C21965. Rare. On
quartzite and limestone.
VERRUCARIA MURALIS Ach. BRY C21966. Rare. On limestone.
XANTHOPARMELIA PLITTIT (Gyelnik ex D. Dietr.) Hale. BRY C21967,
BRY C21968. Rare to common. On moss over quartzite and
quartzite.
XANTHORIA CANDELARIA (L.) Th. Fr. BRY C21969. Rare to common.
On bark of narrowleaf cottonwood.
XANTHORIA ELEGANS (Link) Th. Fr. BRY C21970, BRY C21971, BRY
C21972, BRY C21973. Rare to common. On quartzite, limestone
and moss.
XANTHORIA FALLAX (Hepp in Arnold) Arnold. BRY C21974, BRY
C21975. Abundant. On bark of gambel oak and moss.
XANTHORIA POLYCARPA (Hoffm.) Rieber. BRY C21976, BRY C21977,
BRY C21979, BRY C21981. Abundant. On bark of bigtooth
maple, narrowleaf cottonwood, gambel oak and lignum.
CONCLUSIONS
The lichen flora of Rock Canyon is rich and diverse. The foliose,
squamulose and crustose lichen flora of Rock Canyon is typical of
semiarid montane regions of the Intermountain Area. The
absence of fruticose species is not unusual for Utah. Poor air
quality conditions in Utah Valley are not reflected in the lichen
flora of Rock Canyon. This is likely a result of regular outwash of
air down the canyon to the valley floor.
330
LITERATURE CITED
Anderson, D. C., and S. R. Rushforth. 1976. The cryptogam flora of
desert soil crusts in southern Utah, U. S. A. Nova Hedwigia 28:
691-729.
Flowers, S. 1954. Some lichens of Utah. Proceedings of the Utah
Academy of Science, Arts and Letters. 31: 101-105.
Nash, T. H., and L. L. Sigal. 1981. Preliminary study of the lichens
of Zion National Park. Journal of the Arizona-Nevada
Academy of Science 16: 154-156.
Newberry, C. C., and L. L. St. Clair. 1991. Additions to the Lichen
Flora of Utah I. THE BRYOLOGIST 94: 154-156.
Newberry, C. C. 1991. Lichens of the Uinta Mountains and
Adjacent Intermountain North America. Master's Thesis,
Brigham Young University, Provo, Utah. 236 pp.
Nielsen, H. S. 1960. Lichens of the Wasatch Mountains in central
Utah. Master's Thesis, Brigham Young University, Provo, Utah.
86 pp.
Rushforth, S. R., L. L. St. Clair, J. D. Brotherson and G. T. Nebeker.
1982. Lichen community structure in Zion National Park. THE
BRYOLOGIST 85: 185-192.
St. Clair, L. L. 1984. Lichen distribution along an alpine tundra
ridge in the high Uintas of northeastern Utah, U. S. A. Ph.D.
Thesis, University of Colorado, Boulder. 139 pp.
St. Clair, L. L., S. R. Rushforth and J. D. Brotherson. 1986. The
influence of microhabitat on diversity, distribution and
abundance of corticolous lichens in Zion National Park, Utah
and Navajo National Monument, Arizona. Mycotaxon 26:
253-262.
St. Clair, L. L., and R. B. Warrick. 1987. Acarospora nodulosa
(Duf.) Hue. var. nodulosa: A new record for North America.
THE BRYOLOGIST 90: 48-49.
St. Clair, L. L., C. C. Newberry and G. T. Nebeker. 1991. Catalog of
the lichens of Utah. Mycotaxon 40: 199-264.
St. Clair, L. L., C. N. Newberry, K. H. Hart, and G. T. Nebeker. (in
press) Lichens of Timpanogos Cave National Monument, Utah.
Tuckerman, E. 1871. Lichenes. Pp. 412-413. in S. Watson.
Botany, United States Geological Exploration of the 40th
Parallel.
MY COTAXON
_VolumeL, pp.331—™—~—~————CSCSSantuaary-Maarchh 1994
VALIDATION OF THE NAME
MICROBOTRYUM VINOSUM (USTILAGINALES)
CVETOMIR M. DENCHEV
Institute of Botany, Bulgarian Academy of Sciences
1113 Sofia, Bulgaria
SUMMARY
The fungal name Microbotryum vinosum (L-R. & C. Tul.)
G. Deml & Prillinger was not validly published. The
new combination is thus proposed here. This smut is
reported as new to Bulgaria.
The combination Microbotryum vinosum (L-R. & C.
Tul.) G. Deml & Prillinger in Prillinger, Deml, Doerfler,
Laaser, & Lockau (Bot. Acta 104: 10. 1991) is illegitimate
according to Article 33.2 of the International Code of
Botanical Nomenclature (no reference to the basionym).
Microbotryum vinosum (L.-R. & C. Tul.) Denchev, comb. nov.
Basionym: Ustilago vinosa L-R. & C. Tul.,Ann. Sci. Nat. Bot.,
sér. 3, 7: 96. 1847.
Synonym: Uredo vinosa Berk. in litt. ad Tul. (nom. nud.).
The smut is reported here as new to Bulgaria: mt. Rila,
supra lacum Babreka, 2650 m s.m., 42912’ N, 23°18 E,
22.vii.1909, B. Davidov,SOM 21 017-M. Matrix: Oxyria digyna
(L.) Hill. Spores (6.5-) Mo = 7.840.6 (-10)x (6.5-) Mto = 7.3£0.5
(-9) um (N=100).
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MY COTA XON
Volume L, pp. 333-348 January-March 1994
NEW SPECIES OF GYMNOPILUS
(AGARICALES, CORTINARIACEAE) FROM MEXICO
LAURA GUZMAN-DAVALOS
Instituto de Botanica
Universidad de Guadalajara
Apartado Postal 139
Zapopan, Jalisco 45100
MEXICO
SUMMARY
Six new species of Gymnopilus from Mexico are described, as follow: G.
hemipenetrans from the State of Chiapas, G. commune, G. galerinopsis, G. medius
and G. tuxtlense from the State of Veracruz and G. robustus from the State of Nayarit.
This paper is part of the monograph on Mexican species of Gymnopilus that is
preparing the author.
INTRODUCTION
Continuing the research on the genus Gymnopilus from Mexico, started by the
author seven years ago (Guzmaén-Daévalos & Guzman, 1986, 1991), several specimens
from the Herbaria XAL and IBUG were found that do not fit with the known species,
so they are described as new taxa, before to present the monograph on the Mexican
species that it is in preparation.
Microscopic study was made with 5% KOH, Melzer’s reagent and congo red.
Color of microscopic structures indicated in descriptions was the one observed in KOH.
Omamentation of spore wall is included in spore measures.
334
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Figs. 1-15. 1-5: Gymnopilus commune, |: basidiome, 2: spores, 3: pleurocystidia, 4: cheilocystidia, 5: caulocystidia (all from
Guzman 30481, type). 6: G. galerinopsis, basidiomata (Villarreal 2684, type). 7-12: Gymnopilus galerinopsis, 7: spores, 8:
basidium, 9: pleurocystidia, 10: cheilocystidia, 11: pileocystidia, 12: caulocystidia (all from Villarreal 2684, type). 13-15:
G. hemipenetrans, \3: basidiome, 14: spores, 15: pleurocystidia, type b (all from Guzman 30519, type)(bar: 1, 6 & 13 =
10 mm, 2-5, 7-12 & 14-15 = 4 um).
335
DESCRIPTIONS OF THE TAXA
Gymnopilus commune Guzman-Davalos, sp. nov.
Figs. 1-5
Pileus circa 30 mm diametro, convexo-campanulatus, vel umbonatus, glabrus
ad tenuis fibrillosus, aurantiaco-brunneolus. Lamellae adnatae, subconfertae, angustae,
luteo-aurantiaco. Stipes circa 58 X 5-6 mm, cylindraceus, centralis, pileo concoloro
vel pallidus, velum fibrillosum, zona annular formare. Contextum albidus vel brunneo-
aurantiacus irregulariter. Sporae 6-8.4 X 4-4.8 pum, ellipsoidae, verruculosae,
dextrinoidae. Pleurocystidia 16-28 X 3.2-9.6 wm, ampullacea vel ventricosa,
subcapitata vel capitata, hyalina vel flavida, copiosa. Cheilocystidia 20.8-40 X 4-8 ym,
cylindrico-ventricosa, cylindracea vel ampullacea, capitata vel subcapitata, hyalina vel
flavida. Trama pileus radiata. Pileocystidia nulla. Caulocystidia 16-44.8 X 1.6-5.6 wm,
cylindracea vel cylindrico-ventricosa, capitata vel subcapitata. Fibuligerus. Lignicola
ad Pinus-Quercus sylva, Veracruz, prope Cumbres de Acuitzingo, Puerto del Aire.
Holotypus Guzman 30481 (XAL).
Pileus aprox. 30 mm diam., campanulate convex, with a broad umbo, more
or less glabrous to finelly fibrillose, brownish orange or tanned orange. Lamellae
adnate, subclose, narrow, mustard orange yellow, with a yellowish pruinose edge.
Stipe aprox. 58 X 5-6 mm, equal or ligthly expanded to the base, central, striate-
fibrillose, concolor with the pileus or ligther especially in the apex and in the base,
being yellowish white, solid, with fibrillose remnants from the veil forming an annular
zone. Whitish mycelia in the stipe base. Context whitish to irregularly orange brown,
odour fungoid.
Spores 6-8.4 X 4-4.8 um, ellipsoid, with subthick wall, verruculose, without
germ pore, yellowish brown, dextrinoid. Basidia scarce, aprox. 24.8 X 7.2 um,
tetraspored, sterigmata 3.2 wm long., clavate, hyaline or with yellowish or yellowish
brown content. Pleurocystidia 16-28 X 3,2-9.6 wm, apex 2.4-5.6 wm diam., flask-
shaped or ventricose, with long or short neck, subcapitate or capitate, sometimes non
capitate, hyaline or with yellowish content, very abundant, sometimes clustered.
Cheilocystidia 20.8-40 X 4-8 um, apex 3.2-5.6 um diam., cylindric-ventricose,
cylindrical or flask-shaped, capitate or subcapitate, hyaline or with yellowish content.
Hymenophoral trama subparallel, hyphae with thin wall. Pileus trama radial. Cuticle
with hyphae 2.8-8 um diam., postrate, yellowish brown with incrusted walls.
Pileocystidia absent. Caulocystidia 16-44.8 X 1.6-5.6 wm, apex 3.6-5.2 um diam.,
cylindrical or cylindric-ventricose, capitate or subcapitate, sometimes non capitate,
frequently branched, hyaline or with yellowish content, found only in the stipe apex in
tufts. Clamp connections present. A light yellowish brown pigment is dissolved when
mounted in KOH.
336
HABITAT. Solitary on wood, in a Pinus-Quercus forest.
EXAMINED SPECIMEN. VERACRUZ, Orizaba-Tehuaca4n road, Cumbres de
Acuitzingo, Puerto del Aire (IV Exhibition of Mushrooms from Cérdoba, Ver.), Jul.
24, 1992, Guzman 30481 (XAL, holotype; IBUG, isotype).
OBSERVATIONS. This species is characterized by the glabrous pileus, completly
brownish orange, long stipe, whitish context that changes to orange brown and for
having pleurocystidia clustered, many of them with a narrow and long neck. It is
related with G. subsapineus Hesler, but it is distinguished by the yellowish context,
white stipe, bigger spores, of 7-9(-10) X 4.5-5.5 um from the type study [but 6-8.5 X
4.5-5 um sensu Hesler (1969)] and also bigger pleurocystidia, of 21-38 X 5-8(-12) pm
in type material [25-28(-42) X 5-7 wm sensu Hesler], furthermore the pleurocystidia
do not have long and narrow neck (Hesler, 1969). G. commune is also close with G.
longipes Guzman-Dévalos & Guzman for the stipe size, but the last one has close to
subcrowded lamellae with a long decurrent tooth, spores little wider, inconspicuous
pleurocystidia, shorter and wider cheilocystidia and has pileocystidia (Guzm4n-Davalos
& Guzmdn, 1986). G. liquiritiae (Pers. : Fr.)Karst. has bigger spores and
pleurocystidia subcapitate and inconspicuous (Hesler, 1969; Guzman-Davalos &
Guzman, 1991). The name of the species is due to its common macroscopic habit.
Gymnopilus galerinopsis Guzman-Davalos, sp. nov.
Figs. 6-12
Pileus 30-50 mm diametro, convexus vel convexo-campanulatus, glabrus,
fibrillosus ad areolatus, aurantiaco-rubeolus ad luteo- aurantius. Lamellae sinuatae,
subdistantes, latae, aurantiaco ad aurantiaco-rubeolus. Stipes 35-58 X 5-13 mm,
cylindraceus, centralis, cum pileo concolor vel pallide. Contextum aurantiaco-
rubeolum in pileus, brunneo aurantiacus in stipes. Sporae (6-)6.4-9.6 X 4.4-4.8(-5.6)
um, ellipsoidae, verrucosae, dextrinoidae. Pleurocystidia 25.6-34.4 X 4.4-4.8 um,
tibiiforma ad cylindrico-ventricosa capitata, hyalina, vel brunneo-lutea, infrequens.
Cheilocystidia 28-44 X 3.2-4.8 um, tibiiforma, hyalina vel brunneo- aurantiaca. Trama
pileus radiata. Pileocystidia 1.2-4.8 um diametro, tibiiforma, copiosa. Caulocystidia
2.8-6.4 um diametro, tibiiforma, aliquot cum collo filiform. Fibuligerus. Lignicola ad
Pinus-Abies sylva, Veracruz, prope Cofre de Perote, Los Gallos. Holotypus Villarreal
2684 (XAL).
Pileus 30-50 mm diam., convex to campanulate-convex, glabrous or ligthly
wrinkled, fibrillose or areolate under lens, reddish orange or orange yellow, brick color
when dry. Lamellae sinuate, sometimes with a decurrent tooth, subdistant, broad, light
gon
orange to reddish orange when mature, margin smooth. Stipe 35-58 X 5-13 mm, terete,
broader to the apex and narrower to the base, central, fibrillose, concolor with the
pileus or ligther, base yellowish white, hollow. Veil not seen. Context thick, reddish
orange in pileus, orange brown in stipe and reddish in stipe base. KOH stains dry
pileus and stipe reddish black and context dark vinaceus red.
Spores (6-)6.4-9.6 X 4.4-4.8(-5.6) ym, ellipsoid to ovoid, with subthick wall,
verrucose, without germ pore, yellowish brown, dextrinoid (brown with reddish orange
tinges in Melzer’s reagent). Basidia 26.4-41.6 X 5.6-7.2 um, tetraspored, some
bispored, sterigmata 0.8-5.6 um long., clavate to cylindric-clavate, with or without
central constriction, hyaline or with yellowish or yellowish brown content. Basidioles
very abundant, with yellowish brown or orange yellowish brown content.
Pleurocystidia 25.6-34.4 X 4.4-4.8 um, apex 2.4-6.4 um diam., tibiiform or cylindric-
ventricose capitate, hyaline or with grayish yellow or yellowish brown content, very
rare and incospicuous. Cheilocystidia 28-44 X 3.2-4.8 um, apex 2.4-4.4 wm diam.,
tibiiform, hyaline, or with yellowish, yellowish brown or orange brown content.
Heteromorphous gill edge and fertile. Hymenophoral trama subparallel, hyphae with
thin wall, yellowish. Pileus trama radial, hyphae with thin wall, yellowish. Cuticle with
hyphae 3.2-5.6 um diam., postrate, with yellowish or dark yellowish brown content.
Pileocystidia 1.2-4.8 um diam., apex 2-6.4 um diam., tibiiform, with yellowish or
dark yellowish brown content, abundant. Caulocystidia 2.8-6.4 wm diam., apex 2.6-6.4
pm diam., tibiiform, some with filiform neck, with a cylindrical base prolonged among
the context hyphae, sometimes with subthick wall, especially in the apex and neck,
hyaline or with yellowish or yellowish brown content, present in all stipe long,
including the white base, in palisade in the apex, in tufts or dispersed in the rest,
abundant. Clamp connections present. Lactiferous hyphae present. A yellowish pigment
is dissolved when mounted in KOH.
HABITAT. Gregarious, lignicolous in a Abies-Pinus forest.
EXAMINED SPECIMEN. VERACRUZ, Mpio. of Xico, E zone of Cofre de Perote,
1.5 km N from Ingenio El Rosario, Los Gallos, alt. 2830 m, Oct. 8, 1986, Villarreal
2684 (XAL, holotype; IBUG, isotype).
OBSERVATIONS. G. galerinopsis is defined for having all the cystidia (pleuro-,
cheilo-, pileo- and caulocystidia) tibiiform, to present caulocystidia all the stipe long
and by the pileus color. Because of the tibiiform cystidia it is very close to the genus
Galerina, but this has spores with plage or in case that do not have plage the clamp
connections are absent; furthermore the Galerina basidioma reaction with KOH is
negative (Singer, 1986). By the basidioma color and glabrous pileus, G. galerinopsis
is related with G. oregonensis Murr. and G. picreus (Pers. : Fr.)Karst. sensu auct.,
non Hesler, but it is distinguished by the pleuro- and cheilocystidia flask-shaped or
ventricose, capitate or subcapitate in those species (Hesler, 1969; Bon, 1987; Hoiland,
1990).
338
FO
Figs. 16-28. 16-18: Gymnopilus hemipenetrans, 16: pleurocystidia, type a, 17: pleurocystidia, type c, 18: cheilocystidia (all
from Guzman 30519, type). 19-21: G. medius, 19: basidiome, 20: spores, 21: basidia (all from Garcia-Franco s.n., type).
22-24: Gymnopilus medius, 22: pleurocystidia, 23: cheilocystidia, 24: caulocystidia (all from Garcia-Franco s.n.). 25-26: G.
robustus, 25: spores, 26: basidia (both from Vargas 19, type). 27-28: Gymnopilus robustus, 27: pleurocystidia, 28:
cheilocystidia (all from Vargas 19, type)(bar = 4 um, except in 18 = 5 pm, 19 = 10 mm).
330
Gymnopilus hemipenetrans Guzman-Davalos, sp. nov.
Figs. 13-18
Pileus circa 45 mmdiametro, convexus ad planus, glabrus, aurantiaco-flavidus
ad aurantiaco-brunneus. Lamellae adnatae, confertae, subangustae, aurantiaco-
ferruginae vel brunneo- ferruginae. Stipes circa minus 50 X 5-8 mm, subbulbosus,
subcentralis, apicis subalbidus, pileo concoloro ceterum, velum evanescens. Contextum
flavidum. Sporae 6-8.8 X 4.4-5.6 ym, ellipsoidae vel latus ellipsoideae, verrucosae,
dextrinoidae. Pleurocystidia hyalina vel brunneo-lutea, heteromorphous in tres typus,
a) 20.8-31.2 X 5.2-8.8 um, ampullacea, longicollo, capitata vel subcapitata et copiosa,
b) 17.6-21.6 X 7.2-8 ym, ampullacea, brevicollo, capitata vel subcapitata et
infrequens, et c) 28-36 X 5.2-8.8 ym, ampullacea vel cylindrico-ventricosa et
infrequens. Cheilocystidia 0.8-6.1 um diametro, filiforme, submoniliforme, ventricosa
vel ampullacea, hyalina vel lutea. Trama pileus radiata. Pileocystidia et caulocystidia
nulla. Fibuligerus. Lignicola ad tropicalis sylva, Chiapas, prope Laguna Belgica.
Holotypus Guzman 30519 (XAL).
Pileus aprox. 45 mm diam., convex to convex-plane, glabrous, lightly
fibrillose under lens, dry, yellowish orange to brownish orange. Lamellae adnated,
close, subnarrow, rusty orange to rusty brown. Stipe aprox. 50 X 5-8 mm, terete,
subbulbous, almost central, fibrillose to even, yellowish white in the apex, the rest
concolor with the pileus, hollow, with fibrillose remnants from the veil. Context more
or less thick, yellowish, taste bitter (when dry). KOH stains dry pileus orange brown.
Spores 6-8.8 X 4.4-5.6 um, ellipsoid to widely ellipsoid, with subthick wall,
verrucose, without germ pore, yellowish brown, dextrinoid (discoloring to yellowish
or changing to light reddish brown in Melzer’s reagent). Basidia 20.8-28.8 X 5.6-8.4
pm, tetraspored, some mono-, bi- or trispored, sterigmata 2.4-5.6 um long., clavate,
with or without central constriction, hyaline or with yellowish or yellowish brown
content. Basidioles abundant, hyaline or with yellowish or yellowish brown content.
Pleurocystidia of three types: a) 20.8-31.2 X 5.2-8.8 wm, apex 3.2-4.4 wm diam..,
abundant, gregarious or in small tufts, seldom isolated, flask-shaped, long-necked,
Capitate or subcapitate, sometimes with a thickening in the apex wall, many hyaline,
some with the apex or part yellowish or brown; b) 17.6-21.6 X 7.2-8 um, apex 4.4-5.6
pm diam., solitary, rare, flask-shaped, wide and short-necked, capitate or subcapitate,
hyaline or with yellowish or yellowish brown content, and c) 28-36 X 5.2-8.8 um,
apex 4-6.4 wm, only present very near from the lamellae edge, rare, flask-shaped or
cylindric-ventricose, non capitate, sub- or capitate, hyaline or with yellowish, yellowish
brown or orange brown content, many with a thickening in the apex wall or a content
secretion. Cheilocystidia 0.8-6.1 ~m diam. (only the ends were observed), filiform,
submoniliform, ventricose or flask-shaped, non capitate, sub- or capitate, apex 3.3-5
pm (in sub- and capitate), hyaline or yellowish. Hymenophoral trama subparallel,
hyphae 7.2-31.2 um diam., with thin to subthick wall. Pileus trama radial, hyphae 4.4-
20 wm diam., with thin to subthick wall. Cuticle with hyphae 2.4-6.4 um diam.,
postrate, yellowish, with incrusted walls. Pileocystidia and caulocystidia absent. Clamp
340
connections present. Lactiferous hyphae present. An orange yellow pigment is
dissolved when mounted in KOH.
HABITAT. Solitary on wood causing brown cubical rot, in a perennifolius tropical
forest.
EXAMINED SPECIMEN. CHIAPAS, km 18 Ocozocuautla to Apicpac road, Laguna
Bélgica (Exhibition of Mushrooms from Tuxtla Gutiérrez, Chiapas), Sept. 18, 1992,
Guzman 30519 (XAL, holotype; IBUG, isotype).
OBSERVATIONS. This species is characterized for the three types of pleurocystidia,
the filiform, submoniliform or flask-shaped cheilocystidia, absence of pileo- and
caulocystidia and for its tropical habitat. Macroscopically it is very close to G.
penetrans (Fr. : Fr.)Murr., differing by the adnated to subdecurrent lamellae,
ventricose pleurocystidia, flask-shaped cheilocystidia and the presence of caulocystidia.
Other related species by the pileus color and glabrous surface are G. subsapineus and
G. aurantiophyllus Hesler; from the first one it is distinguished for its emarginate
lamellae with a decurrent tooth, white stipe, cheilocystidia of different form and for the
presence of caulocystidia. G. aurantiophyllus has a olive buff to yellowish context,
lamellae staining rusty orange where bruised, pleuro- and cheilocystidia are of different
form and also has caulocystidia. G. aurantiacus Hesler is near G. hemipenetrans in the
filiform to subtibiiform cheilocystidia, but its pileus is fibrillose-scaly. Other related
species is G. stabilis (Weinm.)Kiihn. & Romagn., presenting spores with plage,
pleurocystidia non capitate or subcapitate and grows in coniferous forest (Kithner &
Romagnesi, 1953 & 1957; Hesler, 1969; Bon, 1987). None of the species mentioned
present the variation in pleurocystidia observed in G. hemipenetrans. The brown
cubical rot observed in this species was cited by Gilbertson (1974) for G. sapineus
(Fr.)Maire.
Gymnopilus medius Guzman-Daévalos, sp. nov.
Figs. 19-24
Pileus circa 80 mm diametro, convexus vel planus, fibrilloso-squamosus vel
fibrilloso-appresus, flavido-brunneus, fibrillula rufobrunnea. Lamellae subdecurrens,
confertae, latae, brunneo- ferruginae. Stipes circa 80 X 8 mm, centralis, fibrillosus,
cremeus, fibrillula brunnea, vellum fibrillosum, zona anullar formare. Contextum
albolutescens. Sporae (7.6-)8-10.4(-11.2) X (4.8-)5.2-6(-6.4) jm, ellipsoidae,
verrucosae. Pleurocystidia 17.6-21.6 X 4-6 um, ampullacea vel cylindrico-ventricosa,
subcapitata vel non capitata, perrara. Cheilocystidia 17.6-24 X 5.2-7.2(-8) wm,
341
ampullacea, cylindracea vel cylindrico-ventricosa, subcapitata vel capitata, hyalina vel
flavida. Trama pileus intricata. Pileocystidia nulla. Caulocystidia 40-64 (seu perlonga)
X 5.6-10.4 ym, cylindracea, non capitata vel capitata. Fibuligerus. Lignicola ad
tropicalis sylva, Veracruz, prope Estacién Bioldgica El Morro de La Mancha.
Holotypus Garcta-Franco s.n. (XAL).
Pileus aprox. 80 mm diam., convex to plane, decurved to straight margin,
surface with appressed or some suberect fibrillose scales, arranged more or less in a
concentric way, dry, brownish yellow to tanned brown, scales dark reddish brown.
Lamellae subdecurrent, close, broad, lamelulla abundant, rusty brown, discoloring in
some places to rusty brownish yellow. Stipe aprox. 80 X 8 mm, equal, base lightly
wider, terete, somewhat flexuose, central, fibrillose, subfleshy to suberose, cream
color, fibrilles light brown, solid, with fibrillose veil remnants forming an annular zone
(not seen in dry material). Context yellowish white, staining yellow when exposed,
odour and taste fungoid. Mycelia white. KOH stains pileus dark amber brown, almost
black.
Spores (7.6-)8-10.4(-11.2) X (4.8-)5.2-6(-6.4) um, ellipsoid, with subthick
wall, verrucose, without germ pore, orange yellowish brown, probably dextrinoid (very
few with reddish tinges in Melzer’s reagent). Basidia 19.2-26.4 X 6-8 um, tetraspored,
sterigmata 0.8-3.2 um long., shortly clavate, with or without central constriction,
hyaline or with grayish yellow content. Basidioles 19.2-23.2(-30.4) X 7.2-10.8 um,
clavate, with dark yellowish brown content. Pleurocystidia 17.6-21.6 X 4-6 wm, apex
3.2-3.6 um diam., flask-shaped or cylindric-ventricose, subcapitate or non capitate,
hyaline, very rare, inconspicuous. Cheilocystidia 17.6-24 X 5.2-7.2(-8) um, apex 4.8-
6.4(-8.8) um diam., flask-shaped, cylindric or cylindric-ventricose, subcapitate or
capitate, hyaline or with light yellowish content, sometimes with brown content in the
apex portion. Hymenophoral trama subparallel. Pileus trama interwoven. Cuticle with
hyphae 3.2-20 um diam., postrate to suberect, join in tufts to form the scales,
yellowish or light brown, with or without incrusted walls. Pileocystidia absent.
Caulocystidia 40-64 (to very long and difficult to measure) X 5.6-10.4 wm, apex 4.4-
7.6 wm diam., cylindric, with or without the end narrowed, apex rounded or capitate,
hyaline, found only in stipe apex in tufts. Clamp connections present. Lactiferous
hyphae not observed. A dark yellowish pigment is dissolved when mounted in KOH.
HABITAT. Solitary, lignicolous, in a subcaducifolius tropical forest.
EXAMINED SPECIMEN. VERACRUZ, Mpio. of Actopan, Biological Station El
Morro de La Mancha, April 19, 1990, Garcia-Franco s.n. (XAL, holotype, IBUG,
isotype).
342
©
30 U 38
37
29
-- -_
-
: Gaga are
s 19, type), 30: G. tuxtlense, pleurocystidia, type b (Guzman
uxtlense, 32: basidioma, 33: basidia, 34:
from Guzman 19830), 37: basidiolum, 38:
diolum, 41: pleurocystidium, type a, 42:
Figs. 29-42. 29: Gymnopilus robustus, pileocystidia (Varga
19671, type). 31: G. robustus, caulocystidia (Vargas 19, type). 32-42: G.
cheilocystidia (all from Guzman 19671, type), 35: basidiomata, 36: spores (both
pleurocystidium, type a, 39: cheilocystidia (all from Guzman 19830), 40: basi
cheilocystidia (all from Guzm4n 19798)(bar = 4 pm, except 32 & 36 = 10 mm).
343
OBSERVATIONS. G. medius is characterized by the appresed fibrillose scaly pileus,
decurrent lamellae and fibrillose remnants from the veil that form a fugacious annular
zone; the pleurocystidia and cheilocystidia are relatively short, the pileus trama is
interwoven and it has caulocystidia. It is close to G. magnus (Peck)Murr., G.
spectabilis (Fr.)A.H.Smith and G. magnificus Guzm4n-Davalos & Guzman; the first
one has a bigger basidioma, pileus fibrillose to somewhat virgate, basidia and
cheilocystidia bigger and pleurocystidia and caulocystidia are absent. G. spectabilis has
a submembranous annulus and basidia, pleurocystidia and cheilocystidia bigger, also
its taste is bitter. G. magnificus has subdistant lamellae, subglobose to ellipsoid spores,
bigger basidia, napiform to piriform cheilocystidia, pileus trama radial and
caulocystidia absent (Hesler, 1969; Mora & Guzman, 1983; Guzmdn-Davalos y
Guzman, 1986).
Gymnopilus robustus Guzman-Davalos, sp. nov.
Figs. 25-29, 31
Pileus 70-90 mm diametro, convexus, plano-convexus vel campanulato-
truncatus, pubescens ad fibrillose-subsquarrosus, subglabrus in aliquot portionis,
exsiccata ochraceus. Lamellae sinuatae, confertae, ventricosae, exsiccata ochracea-
fusca. Stipes 110-130 X 10-18 mm, basis subbulbosis 20-30 mm diametro, centralis ad
excentralis, velum fibrillose, evanescens. Contextum flavidum. Sporae 9.6-11.6 X 7.2-
8.8 um, latus ellipsoidae, subglobosus ad limoniformae, verruculosae, dextrinoidae.
Pleurocystidia 20-29.6 X 6-9.6 um, ampullacea, subcapitata vel non capitata, hyalina
vel lutea, perrara. Cheilocystidia 19.2-28.8 X 5.6-7.2 um, ampullacea vel cylindrico-
ventricosa, capitata vel non capitata, hyalina, infrequens. Trama pileus radiata.
Pileocystidia 30.4-48 X 5.2-8.8 um, cylindracea, lutea. Caulocystidia 29.6-56.8 X 4.4-
8.4 um, cylindracea, cylindrico-clavata, aliquot ventricosa, cylindrico-capitata vel
subcapitata, lutea vel brunneo-lutea. Fibuligerus. Ad palma in tropicalis sylva, Nayarit,
prope Las Varas. Holotypus Vargas 19 (IBUG).
Pileus 70-190 mm diam., convex, convex-plane to campanulate-truncated, with
a Straight or involute edge, pubescent to fibrillose-subsquarrose, some parts
subglabrous, margin smooth, very thin, yellowish ochre when dry, probably rusty
brown when fresh. Lamellae sinuate, with a decurrent tooth, close, ventricose, dark
ochre when dry. Stipe 110-130 X 10-18 mm, more or less equal, base subbulbous of
20-30 mm diam., central to excentric, terete, solid. Veil fibrillose, evanescent, leaving
ochre remnants on stipe apex. Context very thick in the disk, very thin to the margin,
suberose when dry, yellowish. KOH stains dry pileus dark reddish brown.
Spores 9.6-11.6 X 7.2-8.8 um, widely ellipsoid, subglobose to limoniform,
verrucose, yellowish brown, warts imbibed in a well developed perisporium, without
plage and without germ pore, dextrinoid. Basidia 32-36.8 X 7.2-9.6 um, tetraspored,
sterigmata 3.2-5.6 um long., clavate, with central constriction, hyaline. Pleurocystidia
344
20-29.6 X 6-9.6 um, apex 4-5.6 wm diam., flask-shaped, with long or short neck,
subcapitate or non capitate, some with a wall thickening in the apex, hyaline or with
grayish yellow content, very rare. Cheilocystidia 19.2-28.8 X 5.6-7.2 um, apex 2.8-4
pm diam., flask-shaped or cylindric-ventricose, with long or short neck, capitate or non
capitate, many with a wall thickening in the apex, hyaline, rare. Lamellae edge
heteromorphous and fertil. Hymenophoral trama subparallel. Pileus trama radial,
hyphae 4.4-18.4 um diam., thin to thick (1.6 wm) wall, hyaline, yellowish or orange
yellowish. Cuticle very thin, hyphae (3.2-)4.8-11.2 wm diam., postrate, coarsely
incrusted wall with a yellowish, yellowish brown or orange brown pigment.
Pileocystidia 30.4-48 X 5.2-8.8 um, cylindric, apex rounded or attenuate, sometimes
with the medium part inflated, subthick wall, with grayish yellow content, some with
scanty yellowish or yellowish brown pigment incrusted, especially to the base.
Caulocystidia 29.6-56.8 X 4.4-8.4 um, apex 4.4-10.4 um diam. in sub- and capitate,
cylindric to cylindric-clavate, some ventricose with apex rounded or inflated, rare
cylindric-capitate or subcapitate, with subthick wall (0.5-0.8 um), with yellowish or
orange brown vacuolar pigment, some with a yellowish brown pigment incrusted in its
walls, forming a palisade in stipe apex, very abundant. Clamp connections present. A
yellowish pigment is dissolved when mounted in KOH.
HABITAT. Gregarious on a decayed stipe palm, in a caducifolius tropical forest.
EXAMINED SPECIMEN. NAYARIT, Mpio. of Las Varas, Chacala-Chacalilla road,
Nov. 21, 1986, Vargas 19 (IBUG, holotype; XAL, isotype).
OBSERVATIONS. G. robustus is characterized by its big and robust basidioma,
evanescens veil that leaves fibrillose remnants on the stipe, tropical habitat, big and
subglobose to widely ellipsoid spores with perisporium, presence of pleuro-, cheilo-,
pileo- and caulocystidia and by the radial pileus trama. For its basidioma size it is close
to G. magnus, G. spectabilis, G. rugulosus Valenzuela, Guzm4n & Castillo, G.
magnificus and G. subspectabilis Hesler. G. magnus has smaller spores, of 7.5-10 X
5.5-7 um, pleuro-, pileo- and caulocystidia are absent and pileus trama is interwoven
(Hesler, 1969). G. spectabilis also has smaller spores, of 8-10 X 4.5-5.5 um, pileus
trama interwoven and lacks pileocystidia (Hesler, op. cit.). From G. rugulosus is.
distinguished by the membranous annulus, spores of (7.6-)8.4-10.4(-11.2) X 6.4-8(-(.6)
pm, with conspicuous big warts, pileus trama interwoven and pileocystidia absent
(Valenzuela et al., 1981). G. magnificus has smaller spores, of 7.6-9.2 X 5.6-6.4 um
and pileo- and caulocystidia absent (Guzm4n-Davalos & Guzman, 1986). The only
species considered by Hesler (op. cit.) with big spores is G. subspectabilis, of 9-13 X
6-7.5 um, but they are fusiform to sublimoniform, furthermore this species has a pileus
with appresed fibrillose squamules, a submembranous veil and pileocystidia absent.
345
Gymnopilus tuxtlense Guzman-Daévalos, sp. nov.
Figs. 30, 32-51
Pileus 10-25 mm diametro, convexus ad planus, mammiformis ad umbilicatus,
glabrus ad velutinus, hygrophanus, brunneo-aurantiacus vel aurantiaco-ferrugineus ad
flavido exsiccatio. Lamellae adnatae ad sinuatae, subconfertae, latae, brunneo-
ferruginae. Stipes 15-30 X 1-2 mm, centralis, fibrillose byssoideus, pileo concoloro,
basis lanuginosus. Contextum flavidus. Sporae (6.8-)7.2-9.2(-10.4) X 4.4-5.6(-6) wm,
ellipsoidae, verrucosae, dextrinoidae. Pleurocystidia hyalina de duo typus, a) 12-21.6
X 4-7.2(-8) um, cylindrico-ventricosa vel ampullacea, non capitata vel subcapitata,
perrara et incospicua et b) 26.4-56 X 8-16.8 um, ventricoso-rostrata, clavata,
ampullacea, non capitata vel subcapitata, infrequens autem conspicua. Cheilocystidia
14.4-23.2 X (6.4-)7.2-12 wm, ventricoso-rostrata, spatulus-rostrata vel ampullacea,
subcapitata vel capitata, hyalina vel lutea. Trama pileus radiata. Pileocystidia et
caulocystidia nulla. Fibuligerus. Lignicola ad tropicalis sylva, Veracruz, prope
Estacion Biolégica de Los Tuxtlas, UNAM. Holotypus Guzman 19671 (XAL).
Pileus 10-25 mm diam., convex to plane, sub- or mammiform to umbilicate,
glabrous to velutinous, or cottony-fibrillose under lens, half or more than half striate
by transparence, almost to the disk, hygrophanous, orange brown or rusty orange, to
mustard yellow with orange tinges when dry. Lamellae adnate to sinuate, subclose,
broad, rusty brown. Stipe 15-30 X 1-2 mm, equal to lightly inflated to the base or
subbulbous, central, glabrous, cottony-fibrillose under lens, pruinose-fibrillose to the
apex, concolor with the pileus, fibrilles whitish, hollow. Stipe base with a mycelium
halo or a lanose zone, whitish to mustard yellow when dry (probably orange yellow
when fresh). Veil not seen. Context very thin, yellowish. Spore print ochraceous brown
with light tinges orange yellow. KOH stains pileus dark reddish orange brown.
Spores (6.8-)7.2-9.2(-10.4) X 4.4-5.6(-6) um, ellipsoid to amygdaliform, with
subthick wall, verrucose, without germ pore, yellowish brown, dextrinoid (yellowish
with light reddish tinges or brown with reddish orange tinges in Melzer’s reagent).
Basidia (13.6-)16-27.2(-28) X (4.4-)5.6-7.6(-8.8) um, tetraspored, rare bispored,
sterigmata 1.6-4.8 um long., short clavate, with or without central constriction, hyaline
or with yellowish, yellowish brown or orange brown content. Basidioles 16-20.8 X
6.8-9.2 um, spathulate, clavate inflated, spathulate-rostrate, rare or abundant in tufts,
in lamellae edge and faces, hyaline. Pleurocystidia of two types: a) 12-21.6 X 4-7.2(-8)
pm, apex 3.2-5.2 um, cylindric-ventricose, or flask-shaped, sub- or non capitate,
hyaline, very rare and inconspicuous, and b) 26.4-56 X 8-16.8 um, apex (3.2-)4.4-8(-
9.2) wm diam., ventricose-rostrate, clavate, flask-shaped, capitate, subcapitate or non
Capitate, with long or short neck, occasionally with a moniliform neck, sometimes with
an excrescence in the apex, with thin to subthick wall, especially in the apex, hyaline,
rare but conspicuous. Cheilocystidia. 14.4-23.2 X (6.4)7.2-12 um, apex 2.8-5.6 um
diam., ventricose-rostrate, spathulate-rostrate, or flask-shaped sub- or capitate, hyaline
or with yellowish content, some with the apex brown with orange tinges. Lamellae
edge heteromorphous. Hymenophoral trama subparallel. Pileus trama radial, very thin,
346
avenue
Figs. 43-51: Gymnopilus tuxtlense, 43: pleurocystidia, type b (Guzman 19812), 44: pleurocystidium, type a, 45:
pleurocystidia, type b, 46: cheilocystidium (all from Guzman 19798), 47: spores, 48: basidia, 49: basidioles (all from Pérez-
Silva 95, MEXU-19425), 50: pleurocystidia, type b, 51: cheilocystidia (both from Pérez-Silva 95, MEXU-19425)(bar = 4
pm).
347
hyphae with subthick to thick wall. Cuticle with hyphae (2.4-)3.2-13.6 wm diam.,
postrate, few hyaline, many yellowish, some yellowish brown or orange brown, with
or without incrusted pigment. Pileocystidia and caulocystidia absent. Clamp connections
present. Lactiferous hyphae present. No pigment is dissolved in KOH.
HABITAT. Gregarious on branchs or rotten trunks, in a perennifolius tropical forest
and in a coffee plantation.
EXAMINED SPECIMEN. VERACRUZ, Catemaco to Montepio road, Estacién
Biolégica de Los Tuxtlas, UNAM, reserve zone, Jul. 26, 1981, Guzman 19671 (XAL,
holotype; IBUG, isotype); Botanical Garden zone, Jul. 26, 1981, Guzman 19798,
19812 and 19830 (all in XAL); Catemaco region, near to Embotelladora de Coyame,
Nov. 2, 1982, Pérez-Silva 95 (MEXU-19425).
OBSERVATIONS. G. tuxtlense is characterized by the basidioma size, its color when
dry, its tropical habitat and the presence of two types of pleurocystidia. Last
characteristic is only present in G. subbellulus Hesler, sensu Hesler (1969); however
the type of this species (Smith 49838, MICH) has only one type of pleurocystidia (23-
25 X 4-6 um, ventricose). On the other hand, G. hemipenetrans has three types of
pleurocystidia (see under this species). Macroscopically, both G. subbellulus and G.
tuxtlense are close, both have small basidiomata and mustard yellow tinges when dry,
but Hesler’s species grows in coniferous forest, has spores 4.5-6.5 X 3.5-4.5 wm and
presents pileo- and caulocystidia. Other related species is G. radicicola Sing. , separated
by a shorter stipe, smaller spores [6.5-7(-8.5) X 4.5-4.7 um], verruculose to asperulate
and pleurocystidia absent (Singer, 1975).
ACKNOWLEDGEMENTS
The author thanks to Dr. Gast6n Guzman from the Instituto de Ecologia,
Xalapa, Ver. for the critical review of this paper. Dr. Robert L. Shaffer from
University of Michigan Herbarium and Dr. Halling from New York Botanical Garden
kindly permited to review some type materials. Profa. Ma. del Refugio Vazquez from
Guadalajara University finished the ink drawings.
348
LITERATURE CITED
Bon, M., 1987. The Mushrooms and Toadstools of Britain and North-western Europe.
Hodder & Stoughton, London.
Gilbertson, R.L., 1974. Fungi that decay Ponderosa Pine. Univ. Arizona Press,
Tucson.
Guzman-Davalos, L. & G. Guzman, 1986. Hongos del Estado de Jalisco, VII. El
género Gymnopilus (Cortinariaceae). Rev. Mex. Mic. 2: 157-185.
Guzman-Daévalos, L. & G. Guzman, 1991. Additions to the genus Gymnopilus
(Agaricales, Cortinariaceae) from Mexico. Myctotaxon 41: 43-56.
Hesler, L. R., 1969. North American Species of Gymnopilus. Mycological Memoir 3,
Hafner, New York.
Hoiland, K., 1990. The genus Gymnopilus in Norway. Mycotaxon 39: 257-279. |
Kiihner, R. & H. Romagnesi, 1953. Flore Analytique des Champignons Supérieurs
(Agarics, Bolets, Chanterelles). Masson et Cie, Paris.
Kiihner, R. & H. Romagnesi, 1957. VII. Espéces nouvelles, critiques ou rares de
Naucoriacées, Coprinacées et Lepiotacées. Compl. Flore Analytique. Supl.
Bull. Soc. Nat. de’Oyonnax 10-11: 3-94.
Mora, V. y G. Guzman, 1983. Agaricales poco conocidos en el Estado de Morelos.
Bol. Soc. Mex. Mic. 18: 115-139.
Singer, R., 1975. Interesting and new species of Basidiomycetes from Ecuador.
Cramer, Vaduz.
Singer, R., 1986. The Agaricales in Modern Taxonomy. Koeltz, Koenigstein.
Valenzuela, R., G. Guzman & J. Castillo, 1981. Descripciones de especies de
macromicetos poco conocidas en México, con discusiones sobre su ecologia
y distribucién. Bol. Soc. Mex. Mic. 15: 67-120.
MY COTAXON
Volume L, pp. 349-354 January-March 1994
DISCOSIA EUCALYPTICOLA ANAM.-SP. NOV.
T. R. NAG RAJ
Department of Biology, University of Waterloo, Waterloo,
Ontario, Canada. N2L 3G1.
Discosia eucalypticola anam.-sp. nov. Fig. 1
Foliicola. Conidiomata pycnidioidea, hypophylla vel subinde
amphigena, dissita, profunde immersa sed insuper substrato
solummodo area dehiscenti ovale vel circulare procurrenti,
Origine intra-epidermalia vel subepidermalia, lageniformia,
ventere ovale, globoso vel depresse globoso, 80-250 [um Iat.,
60-260 pm alt, et collo late obconico 30-70 [im lat., 40-60 1m
alt., unilocularia, glabra, ut videtur ostiolata, sed per rimam
linearem in telam centralem collarulorum dehiscentia; tela
parietali 10-20 jum cr., ex textura angulari, in stratis extimis e
cellulis crassitunicatis et atrobrunneis, in aream dehiscentiarum
fere nigris, in stratis interioribus pallide brunneis vel subhyalinis
et tenuitunicatis compositus. Conidiophora circum cavitatem
venterem exorientia, cellulae conidiogenae redacta, in muco
involuta. Cellulae conidiogenae lageniformes, rectae vel leniter
curvae, hyalinae, tenuitunicatae, laeviae, 4.5-8 X 2-3 [K = 6.5 X
2.4] jum, usque ad binae annulatim prolificantes. Conidia
fusiformia vel naviculata, 3-septata, hyalina vel subhyalina,
laevia, sine constrictione ad septa, 20-34 X 2.5-3 [kK = 27 X
2.7] um; cellula basalis obconica, basi truncata, 3-4 [X = 3.5] 1m
350
ure. 1. Discosia eucalypticola ex type in DAOM
Fig
eaf showing the
216243. A. Sectional view of a host |
immersed conidiomata. B. v.s. of a conidioma. C.
Conidiogenous cells with developing conidia. D. Mature
conidia.
Gog |
long.; cellulae medianae duae, subcylindraceae, simul 15-26 [x
= 20] um long. (a basi cellula secunda (7-)8-12(-15) [k = 10]
uum long., tertia 8-11(-12) [k = 9.7] um long.); cellula apicalis
conica, apice rotundato, (3-)3.5-4 [x = 3.7] um long.; appendices
filiformes, flexuosae, nonramosae; appendix apicalis singularis,
polaris, 9-11 [k = 10] tm long.; appendix basalis singularis,
subpolaris, excentrica, leniter insuper basi conidiali insertis, 7-10
[xX = 8.5] um long.; ratione conidii long./lat. = 9.8:1.
Foliicolous. Conidiomata pycnidioid, hypophyllous to
occasionally amphigenous, scattered, deeply immersed with only
the oval or circular area of dehiscence visible in surface view,
intra-epidermal to subepidermal in origin, flask-shaped with an
oval, globose or depressed globose venter 80-250 [um wide,
60-260 jum high, and a broadly obconical neck 30-70 [im wide,
40-60 ym high, unilocular, glabrous, seemingly ostiolate, but
dehiscing by a linear split in the central neck tissue; parietal
tissue 10-20 pum thick, of textura angularis, cells thick-walled
and dark brown in the outer layers, almost black in the area of
dehiscence, paler in the inner layers. Conidiophores arising all
around the cavity of the venter, reduced to conidiogenous cells,
colourless, smooth, invested in mucus. Conidiogenous cells
lageniform, straight or somewhat bent, colourless, thin-walled,
smooth, 4.5-8 X 2-3 [k = 65 X 2.4] bm, with 1 or 2
annellations. Conidia fusiform to naviculate, 3-septate, colourless
to almost colourless, wall smooth and without constrictions at
the septa, 20-34 X 2.5-3 [k = 27 X 2.7] tum; basal cell obconic
with a truncate base, 3-4 [k = 3.5] 1m long; 2 median cells
subcylindrical, together 15-26 [x = 20] tm long (second cell
from the base (7-)8-12(-15) [xk = 10] um long, third cell 8-11
(-12) [X = 9.7] uum long); apical cell conical with a rounded
apex, (3-)3.5-4 [x = 3.7] um long; appendages filiform, flexuous,
unbranched; apical appendage single, polar, 9-11 [x = 10] um
352
long; basal appendage single, excentric, inserted slightly above
the conidium base, 7-10 [k = 8.5] um long; mean conidium
length/width ratio = 9.8:1.
Habitat: On leaves of Eucalyptus calophylla and Eucalyptus
sp.
Specimens examined: 1. DAOM 216243 [Holotype], on
leaves of Eucalyptus sp., King’s Park, Perth, West Australia,
25.IX.1992, B.Kendrick; 2. DAOM 216244, on leaves of
Eucalyptus calophylla, John Forrest National Park, West
Australia, 4.X.1992, B.Kendrick.
Known distribution: Australia.
Sutton (1980) published a revised generic description of
Discosia incorporating modern anatomico-ontogenic criteria. The
following characters were considered to be diagnostic: (1)
eustromatic, flattened, black, unilocular to multilocular
conidiomata with 1-many, circular, papillate ostioles; (2) conical
colourless to pale brown conidiogenous cells, and (3) holoblastic,
dorsiventral, straight or slightly curved, 3-4-euseptate, colourless
to pale brown conidia, with (4) two unbranched, filiform, polar
or subpolar appendages one on each of the end cells on the
concave side of the conidia. Subramanian & Reddy (1974),
however, recognized the conidiogenous cells as annellidic,
reported the existence of septate, branched conidiophores in a
few species and stressed the importance of conidium morphology
as well as the relative lengths of conidial cells in differentiating
species. Vanev (1991, 1992a,b, 1993) followed these approaches
when he discussed the species concepts and described three new
species in the genus, although he did not provide details of
conidium ontogeny. Nag Raj (1993) broadened the generic
concept to include superficial and applanate to semi-immersed
or immersed and indeterminate or pycnidioid conidiomata, and
conidium appendages that are branched in at least one species.
353
Species with indeterminate to pycnidioid conidiomata include
Discosia ceanothi (Ramaley) Nag Raj, D. eucalypti (Patouillard)
Nag Raj, D. lauricola Nag Raj, D. novaezelandiae Nag Raj and
D. pyri Koshkelova and form a group to which D. eucalypticola
belongs. Discosia eucalypticola differs from the others in
possessing conidiomata with a well differentiated globose venter
and obconical neck tissue. In conidium morphology it is close to
D. eucalypti, but differs from it in the size of the conidia and the
relative lengths of the individual conidial cells and in the
morphology of its conidiomata. Unlike D. eucalypti it has
reduced conidiophores. The immersed conidiomata with only the
top of the neck tissue visible in surface view, and the almost
black wall tissues of the conidiomata in the venter and neck,
strongly suggest that the fungus is well adapted for life in a
harsh environment. D. eucalypticola appears to be a link in an
evolutionary trend from superficial applanate conidiomata to
immersed, indeterminate to pycnidioid conidiomata.
Acknowledgments
I am grateful to Prof. Bryce Kendrick, Dept. of Biology,
University of Waterloo, and Dr. Amy Rossman, Research
Leader, Systematic Botany and Mycology Laboratory,
U.S.Department of Agriculture, Beltsville, Maryland, U.S.A., for
Critical reviews of the manuscript. Financial support from the
Natural Sciences and Engineering Research Council of Canada
in the form of an operating research grant to Prof. Kendrick is
acknowledged.
References
Nag Raj, T.R. 1993. Coelomycetous anamorphs with appendage-
bearing conidia. 1101 pp. Mycologue Publications, Waterloo,
354
Ontario, Canada.
Subramanian, C.V. and Reddy, K.R.C. 1974. The genus
Discosia. 1. Taxonomy. Kavaka 2: 57-59.
Sutton, B.C. 1980. The coelomycetes, Fungi Imperfecti with
pycnidia, acervuli and stromata. 696 pp. Commonwealth
Mycological Institute, England.
Vanev, S.G. 1991. Species conception and sections delimitation
in the genus Discosia. Mycotaxon 41(2): 387-396.
Vanev, S.G. 1992a. A new species of genus Discosia. Myco-
taxon 43: 69-71.
Vanev, S.G. 1992b. Discosia subramanianii sp. nov. Mycotaxon
44(2): 471-474.
Vanev, S.G. 1993. Discosia arxii sp. nov. Mycotaxon 46: 41-43.
MY COTAXON
Volume L, pp. 355-363 January-March 1994
TRICHOLOSPORUM IN MEXICO AND DESCRIPTION
OF A NEW SPECIES
GASTON GUZMAN
VICTOR M. BANDALA
and
LETICIA MONTOYA
INSTITUTO DE ECOLOGIA
APARTADO POSTAL 63
XALAPA, VERACRUZ 91000
MEXICO
ABSTRACT
Tricholosporum tropicalis Guzman, Bandala & Montoya is
described as a new species from a tropical plantation in the State of
Chiapas, and T. pseudosordidum Sing. is recorded for the first time from
Mexico (State of Veracruz); it was previously known only from Florida
(U.S.A.). A key to the four species of Tricholosporum known from
Mexico is presented. Observations on the type of 7. porphyrophyllum
(Imai) Guzman ex Baroni from Japan are also made, in order differentiate
this species from 7. subporphyrophyllum Guzman.
INTRODUCTION
Tricholosporum Guzman (1975) has been discussed by Baroni
(1982), Mora and Guzman (1983), Guzman et al. (1990) and Bon (1984,
1991). The type species T. goniospermum (Bres.) Guzman ex Baroni was
considered by Singer (1986) in Section Iorigida of Tricholoma, while Bon
(op. cit.) place it in Tricholoma subgenus Tricholosporum (Guzman)
Bon. Seven species of the genus are known to occur in the world
(Europe, North of Africa, Japan, U.S.A. and Mexico), but only two
species, 7. subporphyrophyllum Guzman and T. longicystidiosum
Guzman, Montoya & Bandala are known in Mexico. In this paper two
more species are recorded from this country, one of which is a new
species. Annotations of color are based in Kornerup and Wanscher
(1978).
356
Tricholosporum tropicalis Guzman, Bandala et Montoya, sp. nov.
Figs. 1-4
Pileus circa 58 mm latus, subconvexus vel plano-convexus vel
exigue concavus, glaber, siccus, violaceo-subcaeruleus ad violaceo-
griseolus, vel brunneo-vinaceous vel brunneolus in succus. Lamellae
adnatae, brunneo-incarnatae vel rufobrunneo, vel brunneo-violaceus, acie
concolori. Stipes circa 35 x 8 mm, subbulbosus, glaber, albidus ad
griseo-albidus vel pileo concolor. Caro albida, odore et sapore grato.
Sporae (4-) 4.8-5.6 x 3.2-4 (-4.4) wm, cruciformes, hyalinae,
inamyloideae. Pleurocystidia 30.4-40 (-44) x 5.6-7.2 wm, hyalina,
tenuitunicata, subcylindracea vel subventricosa, rostrato-mucronata.
Cheilocystidia (20-) 24-37.6 (-40) x 5.6-7.2 um, hyalina, tenuitunicata,
pleurocystidiis similia. Cutis gelificata. Trama subregularis. Fibulae
frequentes. Ad terram, solitarius, theobromis cultis. Regio tropicalis.
Mexico, Chiapas, prope Tapachula, A. Jadra s.n. (holotypus XAL).
Pileus approx. 58 mm in diameter, subconvex to convex-applanate
or slightly concave, smooth, dry, somewhat sulcate in dry condition,
bluish violet (18 B5) to grayish violet (18 B4) when fresh to vinaceous-
brown (8D4) to brownish (7D5) in dry, with some _ bluish-violet
(18D3) tinges. Lamellae adnate, + close, pinkish-brown (8B2) to
violaceous-brown (18E5), edges concolorous. Stipe approx. 35 x 8 mm,
central, subbulbous, smooth, white when fresh to gray whitish (6B2 -
6C2) to concolorous with the pileus in dry condition, with irregular
violaceous tinges, base covered with white mycelium. Context whitish.
Odor and taste pleasant. Spores (4-) 4.8-5.6 x 3.2-4 (-4.4) um,
cruciform, hilar appendix conspicuous, thin-walled, hyaline, inamyloid.
Basidia 24-30.4(-32) x 4.8-5.6 ym, claviform, clamped at the base,
hyaline. Pleurocystidia 30.4-40 (-44) x 5.6-7.2 um, abundant, hyaline,
thin-walled, subventricose rostrate and mucronate, sometimes the mucron
as a short cylindrical appendix. Cheilocystidia (20-) 24-37.6 (-40) x 5.6-
7.2 wm, abundant, hyaline, thin-walled, similar to the pleurocystidia in
form; both pleuro and cheilocystidia are projected above hymenial level.
Epicutis as a thin gelatinized layer, hyphae radially arranged, 2.4-6.4 (-8)
yOuyt
Figs. 1-4.- Tricholosporum tropicalis, 1: basidiocarp; 2: spores;
3: pleurocystidia; 4: cheilocystidia (type) (scale bar: 1=2 cm:
2=6 pum; 3-4=10 um) (note: two of the three pleurocystidia
drawing are a little wider than the cheilocystidia, however, the
range of width in both is the same).
358
pm in diameter, thin-walled and smooth. Hymenial trama subregular,
hyphae up to 20 wm in diameter, hyaline and_ thin-walled.
Oleiferoushyphae yellowish or pale yellowish-brown, occasionally in
context and hymenophoral trama. Clamp connections common.
Habitat and studied material. Solitary in soil among branches,
growing in a cacao plantation, in a tropical rain forest region. Mexico,
State of Chiapas, region of Tapachula, Tuxtla Chico, Experimental
Station of Rosario-Iztapa, alt. 50 m, Sept. 25, 1992, A. Jadra s.n.
(holotype, XAL; isotypes ENCB and BR).
Discussion. This tropical fungus is related to the species with
small spores in the genus, such as 7. pseudosordidum (Sing.) Baroni and
T. atroviolaceum (Murr.) Baroni, both from subtropical regions. The
abundant pleurocystidia and cheilocystidia and rather robust basidioma are
the major differences between them. Tricholosporum atroviolaceum lacks
pleurocystidia and cheilocystidia (Baroni, 1982) and 7. pseudosordidum
(see below) has inconspicuous pleurocystidia.
Tricholosporum pseudosordidum (Sing.) Baroni in Mexico
Figs. 6-12
This species was described by Singer (1945) as Tricholoma, from
Florida, U.S.A. and discussed later by Baroni (1982) as Tricholosporum
pseudosordidum. The type of this species, as pointed out by Baroni is
lost. Singer indicated that it was retained in Farlow Herbarium, but the
present authors did not find it, neither there nor at the Herbarium of
Miguel Lillo Institute and University of Florida Herbarium, as indicated
by the respective curators. According to Singer’s original description, the
species apparently lacks pleurocystidia "cystidia none seen; cheilocystidia
(mucronate) and pseudoparaphyses (basidiomorphus) scattered at the edge
of the lamellae, the size of the basidia, very inconspicuous".
In the Gulf area of Mexico (central Veracruz) two specimens were
found, which represent T. pseudosordidum. They have pileus up to 30
mm in diam., plano-convex with a central umbo, pale violaceous (18A3 -
18B4) or yellowish to pale orange (4A3 - 5A4) in the disc and with
violaceous (18A3) or lilaceous (17B2) tinges toward the margin,
hygrophanous. Lamellae adnate, lilaceous or brown-violaceous (18A3 -
18E5). Stipe up to 40 x 4 mm, + attenuated towards the base, whitish to
359
white-vinaceous. The spores are 4.8-5.6 x (3.2-) 4-4.8 (-5.6) um.
Pleurocystidia are very scarce and inconspicuous, (15.2-) 17.6-28 x
(3.2-) 4-5.6 um, hyaline, subcylindric, subfusiform or subventricose,
mucronate or with a short neck. Cheilocystidia 18.4-26.4(-28) x (3.2-)
4-5.6(-6.4) wm, + numerous, but sometimes inconspicuous, similar in
form to pleurocystidia. The basidia are similar in size to both type of
cystidia, (16-) 20-24 (-25.6) x 4.8-5.6 wm, tetrasporic, hyaline to
yellowish, clavate or subcylindric. Epicutis a thin gelatinized layer with
repent hyphae 2.4-5.6 um wide, thin walled. Hymenial trama regular,
with hyphae 1.6-4.8 um wide, hyaline to yellowish and thin walled,
oleiferous hyphae occasionally observed. Clamp connections common.
Singer (1945) described spores (3.7-) 4-5 (-5.5) wm, basidia 24 x 5.3-6
pm and epicutis with irregular filiform, or swollen to clavate hyphae,
which are mostly repent; he did not mention oleiferous hyphae.
The habit of this species is similar to some Lepista in the form
and in the color of the basidioma. As the authors observed the most
typical macroscopic feature of the genus Tricholosporum are the pinkish-
brown to vinaceous brown or violaceous-brown tinges of the basidiomes.
This can also be observed in the color plates of Bon (1984) and Riva
(1988). Singer (1945) described 7. pseudosordidum having white
lamellae but this seems inaccurate, because he considered this species
similar to Lepista sordida (Fr.) Sing., which presents pinkish lamellae as
most of the species in the genus ne and Romagnesi, 1953; Bigelow,
1982, Bon, 1984).
Habitat and studied material. Solitary in subtropical forests with
Quercus and Liguidambar. Mexico, State of Veracruz, I Mushroom
Exposition of Cordoba (specimen collected by Bustos in the road Cordoba
to Huatusco), Aug. 26, 1989, Bandala 2171 (XAL). Road Xalapa to
Huatusco, | km from the deviation to Jalcomulco, Jul. 2, 1983, Chacén
1246 (XAL).
Observations on T. porphyrophyllum and T. subporphyrophyllum
Guzman (1975) based on Tricholoma porphyrophyllum Imai (in
Ito, 1959), described Tricholosporum subporphyrophyllum Guzman by
the presence of pleurocystidia. Guzman and latter Baroni (1982)
considered Imai’s species as Tricholosporum. However, neither Ito
(1959) nor Hongo (1988) indicate either the presence or absence of
hymenial cystidia in their descriptions of T. subporphyrophyllum (as
360
Tricholoma). A revision on the type of T. porphyrophyllum (Imai)
Guzman ex Baroni (Imai, Jul. 31, 1937, SAPA), showed the absence of
pleurocystidia and cheilocystidia. The type presents spores 5.6-6.4 (-7.2)
X (3.2) 4-4.8 (-5.6) um (fig. 5); basidia (18.4-) 20-21.6 x 4-4.8 (-5.6)
pm, tetraspored, hyaline, with cyanophilous or carminophilous bodies
lacking; hymenial cystidia absent; context with cylindric or inflated
hyphae up to 12 um diam., hyaline to yellowish; lamellar trama of
parallel hyphae, 3.2-4.8 ym in diam., frequently with oleiferous hyphae
(yellowish-brown to light amber), and epicutis as a thin gelatinized
layer of + interwoven cylindric hyphae, 1.6-3.2 um wide. This is the
first time that the type of Imai’s species is studied since its description.
KEY TO THE SPECIES OF TRICHOLOSPORUM KNOWN TO
OCCUR IN MEXICO
la. Species with small spores, 4-5.6 um long ............. 2
IbeSpecies' withlargesspores) 15-6 ynvjlongs 22". we 8 ee 3
2a. Pleurocystidia scarce and inconspicuous,
17.6-28 um long. Cheilocystidia +
inconspicuous, 18.4-26.4 pum _ long.
Subtropical species (known only from
the central part of the State of
MV CTacruuz) itineraries. ioctl one ae T. pseudosordidum
2b. Pleurocystidia abundant, 30.4-40 yum long.,
projecting above hymenial level,
Cheilocystidia abundant, 24-37.6 um long.
Tropical species (known only from the
State of Chiapas)
LER? te RE ae ea hte dit Poa ca ads it an caren aero T. tropicalis
3a. Medium size basiomata, pileus 20-60 mm
wide: wustipe 30-50) .uXe0 0-910. mm.
Pleurocystidia 319-37" lum. long:
Cheilocystidia 17-37 ym long. Tropical
species (known only from the States of
Morelos and Oaxaca)
FR ON A tis Rank) ON Orta PAS rel Pa T. subporphyrophyllum
Gee
<a
NY)
~f 13 42
Figs. 5-13.- 5: Tricholosporum porphyrophyllum, spores (type).
6-12: T. pseudosordidum, 6: basidiomata; 7-8: spores; 9-10:
pleurocystidia, 11-12: cheilocystidia (6, 7, 9, 11: Bandala 2171;
8, 10, 12: Chacén 1246). 13: T. longicystidiosum, basidiomata
(type) (scale bar = 10 um, except in6 & 13 = 20 mm).
362
3b. Large size basidiomata, pileus 70-160 mm
wide; stipe 80-130 x 30-50 mm.
Pleurocystidia 30-65 pm_iong.
Cheilocystidia 26.6-48.4 mum __ long.
Subtropical species (known only from the
NE of the State of Morelos)
allie ARR AY PER BONO ey BD SENG Ct G, ant 2 \Citey cae ceanten T. longicystidiosum
(Fig. 13)
ACKNOWLEDGEMENTS
The authors are grateful to Prof. P. Heinemann from Brussels,
who kindly revised critically this paper. They also express their thanks
to the curators of the Herbaria at Farlow (U.S.A.), University of Florida
(U.S.A.), Instituto Miguel Lillo (Argentina) and Sapporo University
(Japan). The help of Fidel Tapia at XAL is appreciated. They are also
indebted to Dr. J. Sanchez from Tapachula who kindly sent the herbarium
specimen of the new species here described.
LITERATURE CITED
Baroni, T.J., 1982. Tricholosporum and notes on Omphaliaster and
Clitocybe. Mycologia 74: 865-871.
Bigelow, H. E., 1982. North American Species of Clitocybe I. Cramer,
Vaduz.
Bon, M., 1984. Les Tricholomes de France et d’ Europe Occidentale. Ed.
Lechevalier, Paris.
Guzman, G., 1975. Un nuevo género y dos nuevas especies de
Agaricaceos mexicanos. Bol. Soc. Mex. Mic. 9: 61-66.
Guzman, G., L. Montoya and V.M. Bandala, 1990. Observations on the
genera Asproinocybe and Tricholosporum, and description of a
new species of Tricholosporum (Agaricales, Tricholomataceae).
Mycotaxon 38: 485-495.
Hongo, T. 1988. On the genus Tricholoma of Japan. Trans. Mycol. Soc.
Japan 29:441-447.
Ito, S., 1959. Mycological flora of Japan, II. Basidiomycetes 5,
Agaricales, Gasteromycets. Yokendo, Tokyo.
Kiihner, R. and H. Romagnesi, 1953. Flore analytique des champignons
supérieurs. Masson, Paris.
363
Mora, V. and G. Guzman, 1983. Agaricales poco conocidos en el Estado
de Morelos. Bol. Soc. Mex. Mic. 18: 115-139.
Riva, A., 1988. Tricholoma (Fr.) Staude. Libreria Editrice Giovanna
Biella, Saronno.
Singer, R., 1945. New and interesting species of Basidiomycetes.
Mycologia 37: 425-439.
Singer, R., 1986. The Agaricales in Modern Taxonomy. Koeltz Sc.
Books, Koenigstein (4th ed.)
ae, i ul
r Tint
oe Ki
ee
MYCOTAXON
Volume L, pp. 365-378 January-March 1994
STUDIES IN THE GENUS PLEUROTUS, Il. THE VARIETIES
OF P. OSTREATUS-COMPLEX BASED IN INTERBREEDING
STRAINS AND IN THE STUDY OF BASIDIOMATA
OBTAINED IN CULTURE
GASTON GUZMAN, LETICIA MONTOYA,
GERARDO MATA and DULCE SALMONES
INSTITUTO DE ECOLOGIA
APARTADO POSTAL 63
XALAPA, VERACRUZ 91000
MEXICO
SUMMARY
The Pleurotus ostreatus-complex ascribed to P. ostreatus (Jacq.
: Fr.) Kumm. var. ostreatus, P. ostreatus var. columbinus (Quél.) Quél.,
and P. pulmonarius (Quél.) Sing. was studied, on basis of the
basidiomata obtained on barley straw in a mushroom farm. The strains
of P. ostreatus var. ostreatus and P. ostreatus var. columbinus are
incompatible, and it is concluded that they are independent species. A
discussion on P. pulmonarius (= P. ostreatus var. florida Eger) and the
misidentifications of P. sajorcaju (Fr.) Sing. are presented. Also a key
to the considered species is included. All these fungi are unknown in their
wild form in Mexico.
INTRODUCTION
Continuing the researches on Pleurotus (Fr.) Kummer by the
authors (Guzman et al., 1991, 1993b), as well as those by the senior
author (Guzman et al., 1980), the interbreeding strains of P. ostreatus
(Jacq. : Fr.) Kumm. complex is discussed, as well as the study of the
basidiomata from strains obtained in culture. The main objective of the
present series, is to define taxonomically and genetically the common and
commercial strains used in Mexico.
MATERIALS AND METHODS
Fourteen strains of Pleurotus ostreatus-complex from different
countries were studied (Table 1). All of them are preserved on malt-agar
366
in the Fungus Strain Collection at Instituto de Ecologia. Concerning with
the origin of the strains, IE-4, 8, 9, 49 and 70 come from INIREB
(Mexico); IE-115 and 140 from Fungi Perfecti at U.S.A.; IE-126 from
Michigan Technological University at Houghton, U.S.A.; IE-129 and 131
from Institute for Fermentation at Osaka, Japan; IE-135, 136 and 137
from Czechoeslovaquia through ICIDCA at Cuba, and IE-138 from
CINVESTAV at Mexico City.
Cultivation of basidiomata at the mushroom farm followed
Guzman et al. (1993a). Basidiomata were obtained at a mushroom farm
on barley straw as substratum. Dilutions of spore prints of strains IE-8,
9, 126, 136 and 140 on distilled water were prepared, in order to produce
monosporic cultures, to which 0.5 ml of this dilution was inoculated on
malt-agar in Petri dishes. About 25 monosporic cultures of each strain
were isolated and 12 of them were selected at random, for determining
mating types. Monosporic cultures of each strain were interbreed,
avoiding reciprocal pairings. Positive pairings were considered when
clamps at joint and at periphery of mycelia were developed, and on the
other hand, negative pairings were considered when clamps were absent.
Monosporic cultures were classified in four types of incompatibility,
according to Eger (1978). Basidiomata were grown under indirect natural
illumination and direct natural illumination, inside and outside the
mushroom farm, in order to observe their color variations. The
microscopic observations were made on slides mounted in 5 % KOH, in
cotton blue in lactophenol, Melzer solution and Congo red. Color in the
descriptions are according to Kornerup and Wanscher (1978). All the
basidiomata studied are deposited at XAL Herbarium.
RESULTS
1) Strains interbreeding
Strains selected according to monosporic cultures, presented a
tetrapolar heterotallic sexual pattern and its respective monosporic
cultures were classified in 4 incompatibility classes, as shown in table 2.
Strains were separated in two groups according to its interbreedin
potential, depending if it was 0 or 100 %. Table 3 shows the
interbreedings, and it can be noted that the strains IE-8 and 126, 9 and
136, 9 and 140, and 136 and 140 were compatible while the rest were
incompatible. In consequence, it can be concluded that P. ostreatus var.
ostreatus is independent of P. ostreatus var. columbinus, considering the
incompatibility.
367
Table 1. Studied strains, its origin and identification
strain species Origin
IE-4 P. pulmonarius Europe (as P. ostreatus
var. florida)
IE-8 P. ostreatus Europe ?
IE-9 P. columbinus Europe (as P. ostreatus)
IE-49 P. ostreatus Europe (as P. sajorcaju)
IE-70 P. columbinus Europe (as P. ostreatus)
IE-115 P, pulmonarius U.S.A. (as P. ostreatus
var. florida)
IE-126 P. ostreatus U.S.A. (as P. ostreatus)
IE-129 P. ostreatus Japan (P. oStreatus)
IE-131 P. ostreatus Japan (P. ostreatus)
IE-136 P. columbinus Europe (as Pleurotus sp.)
IE-137 P. columbinus Europe (as Pleurotus sp.)
IE-138 P. ostreatus Europe (as Pleurotus sp.)
IE-140. P. columbinus U.S.A. (as P. ostreatus)
2) Differences between Pleurotus ostreatus var. ostreatus and P.
ostreatus var. columbinus (Figs. 1-5, 7, 9-13)
Basidiomata from the strains IE-8, 49, 126, 129, 131 and 137
agree with the concept of P. ostreatus var. ostreatus. They have pileus
(25-) 50-100 (-150) mm in diam., hygrophanous, dark grayish-brown
(6C2-6D3) to yellowish-brown (5B3) when dried; sometimes pale grayish-
brown to yelowish-brown (5B3-5B2) or grayish-brown (5B2). Primordia
grayish-brown (5B2, 5C2-6C2) to grayish-white (5SA2), with buff tinges.
Pileus color annotated was observed in basidiomata obtained inside the
mushrooms farm. However, when developed outside the farm under
natural light exposition were darker, dark grayish-brown (7D3) or dark
brown (6D5) to chocolate-brown (6F6); primordia were dark grayish-
brown to chocolate (6F7-6F6), dark chocolate-brown (7F6) to blackish-
brown (6F4) or paler (6D5) at some areas. Lamellae ivory-white to cream
368
color (4A2-4A3), sometimes with pale grayish-brown tinges, decurrent
to subdecurrent. Stipe 10-30 x 7-18 mm or frequently absent, white to
yellowish-white (2A2), excentric, smooth or subvelutinose, villous
towards the base. Context whitish, odor and taste mild or agreeably
fungoid. Spore print grayish-white or white with pinkish tinges (9B2) to
greyish with lilac tinges (14C2), pinkish-grey (14B2), violaceous-grey or
lilac (14C3), to violet with grayish tinges (16D2, 17D3-17D2). Spores
(7.2) 8-10.4 (-11.2) x 3.2-4 (-4.8) um, ellipsoid, hyaline, thin walled.
Basidia 16-28 x 4-6.4 um, claviform, hyaline, tetrasporic. Cystidioid
elements. 16-20.8 x 4.8-5.6 um, rare, ventricose or subcylindric,
mucronate or lecythiform, hyaline, with a mucilaginous drop at apex.
Cystidia absent. Context hyphae 2.4-13.6 um wide, thin walled, rarely
thick walled up to 1.6 um, hyaline, clamped. Hymenial trama iregular;
hyphae 3.2-12 um diam., hyaline, thin or thick walled 0.8-1.6 um,
clamped. Hyphal system monomitic.
Habit pleurotoid, cespitose, but under certain culture conditions
strains IE-126, 129 and 131 frequently developed a clitocyboid habit.
Studied specimens: Guzman 30638; Montoya 1634-A, 2124;
Martinez 288; Salmones 26, 26-A; Tapia 1050 (all from strain IE-8);
Salmones 30 (from strain IE-49); Montoya 1992, 1999 (from strain IE-
126); Guzman 30635, 30636; Montoya 1999-A, 2112 (from strain IE-
129); Montoya 1993, 2000 (from strain 131); Montoya 1905-A (from
strain IE-137).
Kiihner and Romagnesi (1953) considered P. ostreatus "brun. gris,
noiratre ou noir-bleuatre, 5-15 cm. Sp. 8-11 x 3-4 wu". Hilber (1982)
described the pileus of P. ostreatus variable in color, among dark grey,
brownish-buff to violaceous-brown, and spores (5.6-) 9.5- 13.7 x 2.7-3.2-
4.2 um. Bresinsky et al. (1977) reported the pileus dark brown, grayish-
brown or blackish-brown to paler in adult basidiomata. Moser (1983)
considered P. ostreatus with pileus grey, grey-lilaceous, blackish-grey,
steel-grey, grey-brown or blue-blackish, spores 7.5-11 x 3-4 wm and
tramal hypha thin walled.
Basidiomata from the strains IE-9, 70, 135, 136 and 140 agree
well with P. ostreatus var. columbinus, by the dark grayish-blue (22E2-
22E4) to blueish-gray (23C2-23C3) with steel shades, to grayish (23C1),
gray with olivaceous tinges (1B2) or grayish-white with blueish (20B2)
tinges; brownish-buff (5B3) when dried at the disc or over the pileus,
369
Table 2. Incompatibility classes of studied strains
STRAIN INCOMPATIBILITY MONOSPORIC
TYPE CULTURES
IE-8 I
II
III
1
2
4
5
IE-9 I 1
II 8
ITI O76
IV |
TE-126 I 3
II 2)
Ill 6
IV 2
TE-136 I
II
Ill
IV
IE-140 I
II
Ill
IV
7)
©
BO 0760, Oe LZ
, 4, 6, 11
NO Ne WwW NO RY) eRe
‘oN
— 00
=)
—
we)
~
having or not blueish (23C2) tinges or pale yellowish-brown (5B3) to
uniformly grayish-white (5B2), or grayish-brown (6E3) all over, smooth,
sublubricous, but imbricately fibrillose below the cuticle, margin
sometimes translucent striated, (20-) 50-100 (-180) mm in diam.
Primordia 1-8 mm in diam., dark grayish-blue (23D4) to paler (23C1-
23C3), grayish-brown (SE3-5E4, 6E2-6E3, 6D3) to cream-buff (4A3),
sometimes with yellowish (4A2) zones, frequently whitish to grayish,
smooth, lubricous. Lamellae yellowish-white (4A2) to grayish-white
(2A3, 2B2, 2A2), sometimes grayish-green (2B2) to yellowish, sometimes
dichotomic, decurrent, rarely anastamosed at the base. Stipe 15-40 x 3-30
mm, excentric, frequently well developed, yellowish-white (2A2), striated
at first 1/3, subfibrillose, villose at the base. Context whitish, odor and
taste fungoid agreeable. Spore print whitish or white with pinkish (9B2)
370
tinges, grayish-white to lilaceous-gray (14B2), violaceous-gray (14B2-
14C3) to violaceous (17D3-17D2, 16D2). Spores (7.2-) 8-11.2 (-12) x
3.2-4 um, hyaline, thin walled. Basidia 16-25.6 x 4-5.6 (-6.4) um,
claviform, tetrasporic. Cystidioid elements 16-22 x 4-4.8 (-5.6) um,
rare, hyaline, claviform, ventricose or subcylindric, mucronate or
lecythiform, with a mucilaginous drop at apex. Cystidia absent. Context
hypha 2.4-12 (-14.4) um in diam., hyaline, thin walled, some thick
walled up to 2.4 um, clamps numerous. Hymenial trama irregular, hypha
2.4-9.6 um in diam., hyaline, thin walled or some thick walled up to 1.6
um. Hyphal system monomitic.
Table 3. Interbreeding results of studied strains
STRAIN STRAIN STRAIN | STRAIN
IE-126 IE-9 IE-136 IE-140
ee 100 % 0% O % O %
IE- es
STRAIN 100 % 100 %
IE-9
IE-8 & 126: P. ostreatus STRAIN 100 %
IE-9, 136 & 140: P. columbinus IE-136
Habit pleurotoid, caespitose, frequently subclitocyboid due to the
presence of a well developed central stipe. Pileus color varies by the
influence of light, whitish or similar to the colors of P. ostreatus var.
ostreatus; inclusively strains IE-135 and 136 developed the typical blueish
tone only when grown outside the mushroom farm. It was observed also,
that whitish specimens presented a paler spore print.
Studied specimens: Montoya 2089, 2094 (from strain IE-9);
Guzmin 29737, 29738, 30404, 30405, 30406, 30407, 30408, 30411,
30412, 30462; Montoya 1966-A, 1988; Salmones 44 (from strain IE-70);
Montoya 1987, 2102, 2108, 2111 (from strain IE-135); Guzman 30646;
Mata 466, 468; Montoya 1988-A, 1990, 2052, 2062, 2108, 2115, 2119,
2128 (from strain IE-136); Judrez s.n. (13-III-92) (from strain IE-140).
371
Kiihner and Romagnesi (1953) considered the pileus color in P.
columbinus as “gris bleuatre ou verdatre, avec souvent le disque incarnat
ou ocracé 5-9 cm" and spores 10-12 x 3-4 um. Hilber (1982) described
the pileus blue-green, dark tawny or greenish-gray at center and
ochraceous-tawny towards the base and spores 7-10.6-11 x 3.2 -3.5 um.
Bresinky et al. (1977) considered this fungus with the pileus pale steel-
blue or ochraceous at maturity. Moser (1983) considered "Cap grey-blue,
blue-greenish, disc often paler ochraceous or flesh-coloured.... Spores 10-
12 x 3-4 wm". Both P. oStreatus var. ostreatus and P. ostreatus var.
columbinus are very common in Europe, but unknown in Mexico.
3) Pleurotus pulmonarius (= P. ostreatus var. florida) vs. P. ostreatus
(Figs. 6, 8 and 14)
Li and Eger (1979) isolated a strain of Pleurotus from Florida,
U.S.A. which they called Pleurotus "florida" and later as P. ostreatus
"var. florida". However, this name is contaxic with P. pulmonarius,
according to Singer and Harris (1987) and as studied here. P.
pulmonarius was misidentified as P. ostreatus according to several
authors. Eger et al. (1979) mentioned: "the so called P. pulmonarius is
a temperature-tolerant form of P. ostreatus". Many commercial strains
are descendants of Li and Eger strain, as occur with that of Zadrazil
deposited at INIREB (at present Instituto de Ecologia at Xalapa) in 1985
as INIREB-4 (Martinez-Carrera ef al., 1988A), subsequently
misidentified as P. floridanus (Stamets and Chilton, 1983; Guzman-
Davalos et al., 1987) (P. floridanus Singer is an independent species,
probably a Hohenbuehelia). P. ostreatus var. florida Eger is a nomen
nudum (Singer, 1986; Hilber, 1982; Watling and Gregory, 1989). P.
pulmonarius is a whitish fungus like P. djamour (Fr.) Boud., but this
latter species differs in the dimitic hyphal system and cheilocystidia, as
well as in taste and odor (Guzman et al., 1993b). P. pulmonarius seems
common at the Mediterranean zone and in Florida, U.S.A., but is
unknown in Mexico.
The basidiomata produced by strains IE-4, 115 and 138 agree well
with P. pulmonarius, following the concept of Hilber (1982), Bresinsky
et al. (1977) and Moser (1983). The pileus is characteristic in grayish-
brown (5B2-5B3, 5C3-5D3) changing to ivory-white (2A2-4A2), both in
primordia and mature basidiomata, or yellowish when dried,
hygrophanous, smooth, imbricatelly subfibrillose bellow the cuticle,
sublubricous, margin faintly striated, plano-convex to slightly central
372
depressed, at times with a central papilla, (8-) 20-100 (-140) mm in diam.
Lamellae white or whitish (pale 2A2) to ivory-white (1A2-3A2), to
yellowish when dried, decurrent. Stipe (3-) 8-70 (-90) x 2-15 mm,
excentric, striated at first 1/3, fibrillose and villose to hirsute at base,
simple or generally ramified, base more than 30 mm in diam., ivory-
white (1A2, 2A2, 3A2), hygrophanous, compact. Context whitish, odor
and taste fungoid. Spore print white to grayish-white, pinkish-gray (9B2-
14B2) or violaceous-gray (14C3). Spores 7.2-11 (-12) x 3.2-4 ym,
ellipsoid, hyaline. Basidia 20-22.4 x 4.8-5.6 um, claviform, tetrasporic.
Cystidioid elements 20-21.6 x 4-4.8 (-5.6) wm, rare, claviform,
ventricose or subcylindric, mucronate or lecythiform, hyaline, with a
mucilaginous drop at apex. Cystidia absent. Context hypha 4-12 ym in
diam., hyaline, thin walled, at times up to 2.4 um thick. Hymenial trama
irregular, hypha 4.8-10.4 wm in diam., hyaline, thin walled or up to 2.4
pm. Hyphal system monomitic. Hypha clamped.
Habit pleurotoid, but frequently umbelliform, caespitose, ramified
from a common base. Studied material: Guzman 30644, 30645;
Montoya 1903-C, 2012, 2122, 2123, 2125, 2126, 2127, 2130, 2131;
Salmones 32 (from strain IE-4); Guzman 30637, 30639; Montoya 1900,
1902, 2098, 2099, 2100, 2101, 2103 (from strain IE-115); Guzman
30641; Montoya 1901, 2104 (from strain 138).
Specimens grown outside the mushroom farm developed the
characteristic grayish-brown color, while those grown inside were ivory-
white. The specimens of strain IE-138 studied (which are whitish and
stipitate-ramified) have been obtained only inside the mushroom farm,
and according to the thick-walled context hypha, have been considered
tentatively as P. pulmonarius. Recently, Cailleux and Joly (1993)
discussed the differences between P. pulmonarius and P. ostreatus based
in cultures and field observations, and they concluded that they are two
different species growing in France.
4) The misidentifications in Pleurotus sajorcaju
Pleurotus sajorcaju is a paleotropical species very common in
Asia and Africa. Pegler (1977, 1986) considered this fungus as Lentinus
sajorcaju (Fr.) Fr. and characterized it by the infundibuliform pileus and
well developed annulus in the stipe, and dimitic hyphal system. Strain IE-
49 acquired from Europe as P. sajorcaju, as above indicated, produces
typical basidiomata of P. ostreatus var. ostreatus. At present P. sajorcaju
is not known from American Continent.
a73
O00, HAAR. 402,
000 nda DOD
HE ae, 00,
Hl at ( 13 : 14
Figs. 1-14. Microscopic features of the Pleurotus ostreatus-
complex. 1-6: spores, 1-2: 7 ostreatus, 3-5: P. columbinus, 6: P.
pulmonarius. 7-8: basidia, 7: .?. ostreatus, 8: P. pulmonarius. 9-14:
cystidioid elements, 9-11: P. ostreatus, 12-13: P. columbinus, 14:
P. pulmonarius (1: Martinez 288; 2 & 9: Salmones 30; 3 & 12:
Montoya 2089; 4: Montoya 1987; 5: Guzman 30406; 6: Montoya
2012; 7 & 10: Montoya 1999; 8 & 14: Montoya 1905-A; 11:
Montoya 1999-A; 13: Montoya 1990). (Scale bar to spores= 10
pm; Scale bar to basidia and cystidioid elements = 12 pm).
374
DISCUSSION AND CONCLUSIONS
Considering the incompatibility during interbreeding of European
and North American strains of P. ostreatus var. ostreatus and P.
ostreatus var. columbinus, it can be concluded that P. ostreatus var.
columbinus is an independent species, so it must be considered as:
Pleurotus columbinus Quél., apud. Bres., Fungi Trid. 1, p. 10,
1881
Synonyms:
" P. columbinus Bres., Fungi Trid. 1, tab. VI, 1881"
P. ostreatus var. columbinus (Quél.) Pildt, Champ.
Europe, p. 120, 1935
P. ostreatus var. columbinus (Quél.) Quél. s. auct.
Eger et al. (1979) considered P. columbinus as a simple "form"
of P. ostreatus, with blue pileus. Certainly, under culture conditions the
basidiomata of P. columbinus frequently lose the characteristic blue
colors, but under natural conditions or when grown outside the mushroom
farm, the pileus always developes blue colors. According to the
bibliography, P. columbinus probably is more frequent in coniferous
woods and P. ostreatus in deciduous woods. P. pulmonarius is the correct
epithet to the commercial strain known as "Pleurotus ostreatus var.
florida" or “Pleurotus florida", actually both nomina nuda. Pleurotus
floridanus (Stamets and Chilton, 1983) is obviously a misidentification.
P. sajorcaju is unknown in American Continent, and some commercial
strains of P. ostreatus have been misidentified with that paleotropical
species.
As for cystidioid elements observed in the three studied species,
these elements are of no taxonomic value and agreed well with structures
described by Hilber (1982) in P. ostreatus var. ostreatus and P. ostreatus
var. columbinus as " unfunctional basidia, supporting conidia ". Hilber
considered these elements erroneously as caulocystidia, but they are born
from lamellae extended over the stipe, as he shows in the illustrations
(eg. Fig. 124) where a reduced hymenium is represented. This author
also called these structures as "reduced hymenial elements like,
supporting globose conidia, 2.6 um in diam., with ability of reproduce
asexually", but in figure 135 he shows mucilaginous drops sheathing the
apex, aS the writers observed in the studied materials. Probably the
cystidioid elements could, in fact be aberrant basidia produced under
culture conditions, as observed in anamorphs of Pleurotus smithii
(Guzman et al., 1980, fig. 5).
375
In relation to basidiomata color, light affects significantly on its
variation. Bresinky et al. (1977) observed that differences between P.
ostreatus and P. columbinus on basidiomata color, when grown at low
temperatures and high intensities of light, appear whitish. P. ostreatus
var. ostreatus grown at the mushroom farm, either inside under indirect
natural ilumination or outside under direct natural ilumination, presented
different colors; it was observed that the characteristic grey, lilaceous-
gray, greyish-brown or blackish colors of pileus, vanished inside the
mushroom farm, while those outside developed the characteristic colors.
P. columbinus grown outside the mushroom farm presented more intense
blue colors than those grown inside, these latter were pale brownish-buff,
similar to those of P. ostreatus, while P. pulmonarius is whitish or ivory-
white when grown inside, but pale brownish-buff under natural
illumination.
Basidiomata form tend to vary also under culture conditions. The
typical form in studied species is shell-like. However, under poor
aeration, high concentrations of CO, and bad illumination, stipitate forms,
with infundibuliform pileus and deeply decurrent lamellae are developed,
so the basidiomata form is not a good taxonomic character on species
diagnosis in Pleurotus, but most basidiomata of P. pulmonarius showed
a tendency to have a well developed and ramified stipe, not observed in
P. ostreatus neither on P. columbinus, although this later tends to present
+ well developed stipes, but never ramified. The stipitate ramified form
of P. pulmonarius is somewhat like P. cornucopiae Paul. : Fr. (= P.
cornucopioides Pers.), a species known poorly in the American
Continent.
Key to species of Pleurotus ostreatus- complex
la. Pileus dark grayish-blue,
grayish-blue or grayish to
steel grayish. Spores 8-11.2
(-12) um length. Hyphae
thick walled, up to 2.4 wm.
Common in_ coniferous
forests in Europe and North
America (unknown in
RC NACO NU ADE WR ue ah yi Mave) Congas ott. P. columbinus
WELLES NATITOUIL DELITS IV CLQITICS i tal onsh ste Ce esc AN EU A MOAT No Ink 2
376
2a. Pileus dark grayish-brown to
pale grayish brown. Spores
8-10.4 (-11.2) pm_ long.
Hyphae thin walled, up to
Lor pm Common! in
deciduous temperate forests
in Europe and North
America (unknown in
Mexico) 27 PAS aR, ceca ee ee Teak, Oe P. ostreatus
2b. Pileus pale grayish-brown to
ivory-white.Spores 7.2-11 (-12)
pm long. Hyphae thick walled,
up to 2.4 yum. Common in
subtropical or deciduous forests |
(noOtan’Mexico) see Lee eee bee ee, Meer Sane da P, pulmonarius
ACKNOWLEDGMENTS
The authors express their acknowledgements to Michigan
Technological University (U.S.A.), Institute for Fermentation at Osaka
and ICIDCA at Cuba for the exchange of strains. They are grateful to R.
de Leon Chocooj from Guatemala, F. Voguel from Fortin (Mexico) and
C. Dickel from Coatepec (Mexico), who kindly donated some strains. It
is also appreciated the collaboration of V. Alvarez, R. Pérez, M. Juarez,
R. Medel, F. Tapia, M. E. Ramirez and J. Lara from Instituto de
Ecologia, for their valuable help at the herbarium, laboratory,
computation and mushroom farm. They are gratefull to V. Bandala who
kindly prepared the line drawings and critically revised this paper. Dr. D.
N. Pegler from Kew, kindly revised this paper. B. Webel helpfully
translated information from German.
LITERATURE CITED
Bresinsky, A., O. Hilber and H.P. Molitoris, 1977. The genus Pleurotus
as aid for understanding the concept of species in Basidiomycetes. In:
Clemencon, H., The species concept in Hymenomycetes. Cramer,
Vaduz.
Cailleux, R. and P. Joly, 1993. Pleurotus ostreatus (Jacq. : Fr.) Kummer
et P. pulmonarius (Fr.) Quél. : etudes préliminaires. Bull. Soc.
Mycol. France 109: 27-41.
377
Eger, G., 1978. Biology and breeding Pleurotus. In : Chang, S.T. and
W.A. Hayes (eds.), The Biology and Cultivation of Edible
Mushrooms. Academic Press, New York.
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378
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MY COTAXON
Volume L, pp. 379-407 January-March 1994
OMPHALINA SENSU LATO IN NORTH AMERICA 1-2.
1: OMPHALINA WYNNIAE AND THE GENUS CHRYSOMPHALINA
2: OMPHALINA SENSU BIGELOW
LORELEI L. NORVELL, *SCOTT A. REDHEAD & JOSEPH F. AMMIRATI
Department of Botany KB-15, University of Washington,
Seattle, Washington 98195 USA;
*Centre for Land & Biological Resources Research, Research Branch,
Agriculture & Agri-Food Canada, Ottawa, Ontario CAN KIA OC6
ABSTRACT: The four year observation of an Oregon population of Omphalina wynniae has
facilitated reevaluation of its generic placement and that of other North American omphalinoid
agarics. In Part 1, the species is illustrated and described in detail, its nomenclatural history
discussed, and the new combination Chrysomphalina grossula proposed for it. The
genus Chrysomphalina is reevaluated: new combinations C. chrysophylla var.
salmonispora and C. chrysophylla var. hoffmanii are made; Gerronema_ strombodes
is removed from Chrysomphalina and the new combination G. xanthophyllum is made for
a vicariant European taxon which has been confused with G. strombodes. A key to the known
species and varieties of Chrysomphalina is presented. In Part 2, the division of Omphalina
sensu Bigelow and the generic concepts of Gerronema, Haasiella, Omphalina, Phytoconis,
Pseudoarmillariella, and Rickenella are discussed. New combinations are Omphalina
hohensis and O. marchantiae, and Hygrocybe luteo-omphaloides nom. nov.
is proposed for O. occidentalis. A key to generic segregates of Omphalina sensu Bigelow and
expanded generic descriptions are provided.
KEY WORDS: Basidiomycetes, Agaricales, Tricholomataceae, Camarophyllus,
Chrysomphalina, Clitocybe, Gerronema, Haasiella, Hygrocybe, Omphalina, Phytoconis,
Pseudoarmillariella, Rickenella, Pacific Northwest, nomenclature, new combinations, generic
key, anatomy, ecology, morphology
Introduction
Since Bigelow's 1970 monograph of the genus Omphalina in North
America, several revised classifications of the species he treated have been
published. One of the species, O. wynniae (Berk. & Br.) Ito, is rare, and on
this continent is geographically restricted in distribution. Fresh material was
not available to Bigelow; however, the availability of living material in Oregon
has allowed us to study the species and its generic placement in detail (Part 1),
which in turn has prompted reconsideration of the generic concepts of other
North American omphalinoid agarics (Part 2).
380
Methods: Microscopical data were taken either from fresh material,
observed in water, or dried basidiomes, rehydrated and mounted in Melzer's
reagent or 3% KOH. Capitalized color notations are from Ridgway (1912).
Collections cited under ‘SPECIMENS EXAMINED’ are deposited in the
University of Washington Herbarium (WTU) unless otherwise stated.
Herbarium abbreviations are from Holmgren et al. (1990). Norvell collection
numbers start with initials, year, month, and day, i.e. LLN 92.10.28-3 = Oct.
28, 1992, 3rd collection.
Part 1. OMPHALINA WYNNIAE AND THE GENUS CHRYSOMPHALINA
Weekly monitoring of an Oregon population of O. wynniae from
1990 to 1993 facilitated reevaluation of the generic placement of the species,
with macroscopic and microscopic features, pigmentation, and biology all
supporting its placement into the segregate genus, Chrysomphalina Clg.
Additionally, a review of the literature revealed the existence of an earlier
name, Agaricus grossulus Persoon, for the taxon.
A detailed consideration of generic placement follows a complete
species description and nomenclatural discussion.
CHRYSOMPHALINA GROSSULA (Pers.) comb. nov.
Figs. 1-7
Basionym: Agaricus grossulus Pers. Mycol. Eur. 3: 110. 1828.
= Omphalina grossula (Pers.) Sing. Persoonia 2(1): 29. 1961.
= Camarophyllus grossulus (Pers.) Clg. Beih. Z. Mykol. 4: 55. 1982.
= Gerronema grossulum (Pers.) Sing. Beih. Sydowia 7: 14. 1973.
= Agaricus umbelliferus var. abiegnus Berk. & Br. Ann. Mag. Nat. Hist. Ser. 4,
XV: 28, #1143. 1875.
=Omphalia abiegna (Berk. & Br.) J. Lange. Dansk. Bot. Arkiv 6: 13.
1930:
= Omphalina abiegna (Berk. & Br.) Sing. Lilloa 22: 212. 1951.
= Hygrophorus wynniae Berk. & Br. Ann. Mag. Nat. Hist. Ser. 5, III: 208, #1781.
1879.
= Omphalia wynniae (Berk. & Br.) Quél. C.R. Assog. Franc. Av. Sci. (La
Rochelle, 1882) p. 390. 1883.
= Omphalina wynniae (Berk. & Br.) Ito. Myc. Fl. Japan 5(2): 128. 1959.
= Omphalina bibula Quél. Ench. Fung. p. 44. 1886.
= Omphalia bibula (Quél.) Sacc. Syll. Fung. 5: 324. 1887.
=Omphalina umbellifera var. citrina Quél. Ench. Fung. p. 44. 1886.
= Omphalia bibula var. citricolor Rolland. Bull. Soc. Myc. France 7: 94. 1891.
= Omphalia bibula f. citricolor (Rolland) Konr. & Maubl. Icon.
Sel. Fung. p. 288. 1934.
381
Fig. 1. Chrysomphalina grossula (LLN 92.10.25-4) in situ.
Approximately actual size.
Selected descriptions and illustrations: NORTH AMERICA: Bigelow (1970, 25-
26, Fig. 10 as Omphalina wynniae); Phillips (1991, 76-77 as Omphalina wynniae); Smith
(1937, 481 as Omphalia abiegna); EUROPE: Breitenbach & Kranzlin (1991, 100-101 as
Camarophyllus grossulus); Clémengon (1982, 58 as Camarophyllus grossulus); Cooke (1886.
Pl. 905 [911], Fig. A as Hygrophorus (Hygrocybe) wynniae); Josserand (1955: 74-76 as
Omphalia abiegna); Konrad & Maublanc (1933: Pl. 235, Fig. 2 as Omphalia wynniae); Kihner
& Romagnesi (1953, 123 as Omphalia abiegna); Moser (1983, 98 as Omphalina grossula);
Persoon (1828,110, Tab XX VI Fig. 6 (NOT Fig. 2) as Agaricus grossulus); Quélet (1886, 44-45
as Omphalina umbellifera var. citrina); Rolland (1891, 94-95 Pl. VI, Fig. 1 as Omphalia bibula
var. citricolor); Ryman & Holmasen (1984, 260 as Omphalina grossula); AFRICA: Malencon &
Bertault (1975: 202-204 as Omphalina abiegna); ASIA: Ito (1959, 128 as Omphalina
wynniae); Kawamura (1929, No. 58 & 1954, 314 (Fig. 311) both as Hygrophorus wynniae).
PILEUS: 2-35 (60) mm, convex to plano-convex with incurved
margin when young, becoming convexo-umbilicate to uplifted with age;
extreme margin even, crenate or occasionally sulcate; surface moist,
hygrophanous, striate from disc to extreme margin, glabrous; color of disc
and striations initially yellow to brownish yellow or greenish yellow (Wax
Yellow, Citron Yellow, Old Gold, Olive Lake), with age becoming paler
(Citrine, Old Gold, Olive Lake) or even whitish, color of margin yellow to
greenish yellow (Wax Yellow, Massicot Yellow, Citron Yellow, Pyrite Yellow),
with extreme age entire cap almost white; context thin (1 mm over gills), pale
yellow (near Massicot Yellow). LAMELLAE: strongly decurrent, initially
ending at the same point on stipe apex, distant (L + 1 ca. 9/cm at edge and
382
5/cm at midpoint), arcuate, 1-1.5 (3) mm broad, 0.5-0.8 (1.0) mm thick; often
intervenose, lamellulae 1- or 2-tiered, regularly interspersed; edges even; color
generally yellow to greenish-yellow (Marguerite Yellow, Citron Yellow, Pyrite
Yellow, Sulphur Yellow) becoming slightly paler to whitish on exposure or
with age. STIPE: central, 5-40 (55) mm long, more or less equal 1.5-7 (10)
mm at apex, evelate, usually hollow; surface appearing glabrous to the naked
eye, appearing sparsely minutely pubescent under magnification; color
overall yellow or greenish-yellow (Massicot Yellow, Citron Yellow) gradually
paling toward base with age; context pliable, pale yellow (Massicot Yellow).
Odor and taste not distinctive. SPORE PRINT whitish in thin deposits.
SPORES: 5.9 - 9.6 X 3.7-5.5 (6) um, ellipsoid to subellipsoid with
conspicuous obtuse apiculus (Fig. 3); hyaline, smooth, thin-walled, inamyloid,
frequently containing one large central oil droplet when fresh and viewed in
water or KOH [dried spores older than ten years contain many small droplets].
BASIDIA: 33-48 X 5-8 Um, 4-spored (rarely 2-spored), cylindrical to
narrowly clavate with sterigmata ranging in length from 3 - 7.4 (10) um;
packed with the basidioles into a dense hymenial layer (Fig. 4); ratio of
basidioles to sterigmate basidia very high, HYMENIAL CYSTIDIA: absent.
OLEIFEROUS HYPHAE: _ flexuous, with dense, yellowish refractive
contents, 4-6 tum diam, occasionally present in all tramal tissues and in the
stipitipellis. CLAMP CONNECTIONS: absent throughout the basidiome.
PILEIPELLIS: in radial section a cutis of compact parallel to
subparallel, broadly cylindrical, repent, frequently septate hyphae (3)-6-13
uum diam; (occasionally branching or fusing with other elements in age);
occasional cylindrical to tapered cystidioid elements with rounded apices
exserted from the surface (Fig. 6); hyphal walls thin, smooth, neither
gelatinized nor with incrusted pigments, and with slight banding from
presence of intraparietal pigment; faint brownish-yellow intracellular pigments
present when viewed in water. PILEAL TRAMA: a loosely radiating mesh-
like structure of septate, hyaline, smooth, thin-walled, nongelatinized hyphae,
8-20 (22) tum diam, often ampullaceous at septa, giving many cells a femur
(bone-like) shape (Fig. 5).
HYMENOPHORAL TRAMA: mature lamellae in cross section with
a relatively wide mediostratum of greatly inflated cells similar to those in the
pileal trama bracketed by a thin layer of narrow descending hyphae giving
rise to the subhymenium, the latter of perpendicular hyphae branched in a
candelabra-like manner which give rise to the basidia (Fig. 4). [In young
lamellae the mediostratum is composed of parallel to subparallel hyphae (the
parallel alignment being more noticeable in the central portion) which
become highly inflated (>12 sm diam) in older lamellae; hyphal walls
smooth, thin, neither gelatinized nor incrusted; yellowish intracellular pigment
present throughout; bracketing subhymenium of 1-4 individual strands of
parallel, narrow, cylindrical, septate hyphae 10 um diam, with age becoming
more sinuous, less regular and obscured by inflated hyphae of the
mediostratum (often visible only in squash mounts); perpendicular to these
strands, a layer of irregularly shaped di-/ trichotomously branched inflated
hyphal elements, 3-5 tm diam, each giving rise to 2-3 basidioles, basidia, or
383
other subhymenial elements; this candelabra-like layer, 20 Um wide in young
specimens, with basidia arising from different levels.]
STIPITIPELLIS: a cutis of repent, subparallel, somewhat inflated,
septate hyphae 3.7 - 12.5 (17) um diam, with occasional exserted cystidioid
elements (Fig. 7); pale yellowish intracellular pigment present; walls thin,
smooth, neither gelatinized nor pigment-incrusted. STIPITITRAMA:
composed of inflated, septate, hyaline, thin-walled vesicular hyphae 8-22 um
diam.
ECOLOGY AND DISTRIBUTION: gregarious to caespitose on
water-soaked coniferous wood, bark chips, debris (occasionally found on
angiosperm wood mixed with colonized coniferous wood) in mixed forests or
parks at elevations from 30 m to 1100 m. Chrysomphalina grossula (as
Omphalia abiegna) was first reported in North America from Washington
state (Smith, 1937). In North America this species is uncommon, restricted to
the Tsuga heterophylla/Pseudotsuga menziesii zone of the Pacific Northwest
where it is normally found in small numbers on coniferous debris. A report
from Alberta by Schalkwyk-Barendsen (1991) was based on a collection of
Clitocybe, sect. Candicantes [specimen in DAOM examined].
Fig. 2. Young, more darkly colored Chrysomphalina grossula basidiomes
exhibiting hollow stipe (LLN 90.10.20-2). Slightly larger than natural size.
384
NORTH AMERICAN SPECIMENS EXAMINED: CALIFORNIA:
Humboldt Co, Arcata -- X//.18.1986 BLT 606 (HSC). IDAHO: Bonner Co,
Nordman -- X.6.1956 AHSmith 54126 (MICH); X.8.1956 AHSmith 54271 (MICH);
X.11.1956 AHSmith 54452 (MICH); [X.23.1964, AHSmith 70705 (MICH); Priest
Lake -- /X.10.1966 RWatling 2892 (E); /X.29.1968 AHSmith 76757 coll.
RHPetersen (MICH); Shoshone Co -- /X.21.1966 OKMiller 3351 (E). OREGON:
Clackamas Co, Mt. Hood -- X.3.1946 AHSmith 24026 (MICH); Lincoln Co,
Van Duzer State Park -- LLN 92.11.10-2; Multnomah Co, Portland -- LLN
90.10.20-2; LLN 90.10.28-1,2,3,4,5; LLN 90.11.26-3; LLN 91.04.07-1,2; LLN
91.05.12-1; LLN 91.11.19-1 (DAOM); LLN 91.11.23a-c; LLN 92.01.02-1,2 coll.
OLNorvell 3 (DLamoure pers. herb.); LLN 91.01.12-1; LLN 92.01.25-1; LLN
92.01.31-2; LLN 92.02.07-1,2,3,4,5,6; LLN 92.02.29-1; LLN 92.04.19-1; LLN
92.04.28-3,4; LLN 92.05.02; LLN 92.05.03-3; LLN 92.11.10-1 coll. GTNorvell;
LLN 93.04.11; LLN 93.04.18-1,2,3,4,5; LLN 93.04.25-1,2. WASHINGTON:
Clallam Co, Lake Crescent -- X.9.1935 AHSmith 3059 (MICH); X./3.1935
AHSmith 3123 (MICH); Columbia Co, Tucannon River -- /X.22.1984 JFAmmirati
9014; King Co, Seattle --LLN 90.11.26-4 coll. DVBark; LLN 90.12.11-2 coll.
DVBark; Lewis Co, Longmire -- V///.1.1948 AHSmith & DEStuntz; Pierce Co,
Green Lake -- X.2.1]952 AHSmith 40415 (MICH); X./9.7952 Stuntz 7554 AHSmith;
Mt. Rainier Natl Park, Lower Tahoma Creek -- X.22.1972 JWLennox 1412;
Whatcom Co, Mt. Baker National Forest, Noisy Creek Preserve -- LLN 92.03.28-1
coll. JFAmmirati.
EXTRALIMITAL SPECIMENS EXAMINED: CZECHOSLOVAKIA: Mt.
Boubin Nature Reservation -- /X.28.1970 RSinger C5205 (F). SCOTLAND:
Hanzyll, Ben More Garden -- /X.6.1963 DMHenderson 7087 (E); Herefordshire,
Covenhope -- X/.21.1959 PDOrton 2062 (E); Invernesshire, Cullodon Visitor
Center -- X.20.1989 RWatling 21679 (E); Keltshire, Black Craig --V//.25.1970
RWatling 7618 (E); Lothenchurch, Lachane -- /X./3.1957 RWatling 37C (E);
Perthshire, Rannoch, Dall -- X.7.1976 PDOrton 4855 (E); Straloch --
VIII.18.1978 RWatling 12844 (E); Skye, Treaslane -- X.21.1980 AP & FM Bennell
NG 394524 (E). SWEDEN: Lena Parish, Storveta --/X.28.1937: SLundell ex
Fungi Exsicc. Suec #517 (BPI); SLundell, det. RSinger 5563 (F); Smaland, Nassjo
Parish, [Stuntamalen] -- X.9.1945 G Haglund 572 (MICH); SWITZERLAND:
Maschwanden/ZH Meggenwald -- X.20.1975 FKranzlin 2010-75 K1 (LU);
Obfelden/Oberholz --X//. 11.1988 JBachler 1112-88BA1 (LU).
Observations and Notes
In 1981 brilliant yellow basidiomes of Chrysomphalina grossula were
noticed in a forested garden on the outskirts of Portland, Oregon. In
September of 1990, the species reappeared. A few basidiomes were found
approximately 150 feet away from the original site on a trail leading into a
second-growth forest. In both instances the fungus occurred on chipped bark
imported to the garden for use as mulch. During the first few months
following its reappearance, the current population expanded from its original
location along the path for approximately fifteen feet. Subsequently in the
summer of 1991 basidiomes extended over 150 feet in large numbers over the
entire length of the trail; spread of the original population, now extending
over a distance of 700 feet, has continued to the present time.
Observations of individual basidiomes over time have revealed that this
species exhibits considerable color variation throughout its development. The
basidiomes are long-lived; some specimens have been observed in situ for
385
over a month. The greatest intensity of color, a brilliant greenish-yellow,
occurs upon emergence from the substrate. Throughout this period the pileus
and stipe base gradually lose pigments. Intense coloration is lost from the
more exposed areas; the pilei tend to fade from an intense olive-yellow
through greenish-brown or tan to almost white, while more protected areas of
the basidiomes, such as the lamellae and stipe apex, retain an intense greenish-
yellow hue for a much longer time. Eventually the entire basidiome can
become pallid. This developmental color variation is paralleled by differences
in innate pigmentation: instances of partial albinism, where yellow pigments
are missing from the onset, also have been observed in the Oregon population
(LLN 90.10.28-2, LLN 91.11.23c, LLN 93.04.18-1&2.). This variability of
the intrinsic color found in different forms of C. grossula was noted by
Josserand (1955), who also commented on the occurrence of a double-
pigment system: one citrine, the other a bister-fuscous.
The lamellae thicken considerably over the life of the basidiome; as a
result the lamellar trama varies from subregular to irregular depending upon
the disruption caused by inflation of the hyphae in the mediostratum (Fig. 5).
Simultaneously the hymenium thickens slightly. At any given time there are
few sterigmate basidia relative to basidioles (Fig.4); this is due to the continual
rejuvenation of the hymenium over the extended life-span of the basidiome.
The range in size of the individual basidia in C. grossula is similar to
that observed in C. chrysophylla (Fr.) Clg. In his discussion of subhymenial
and basidial development in that fungus, Kiihner (1928) remarked "each cell
of the subhymenium gives rise to an ascending crooked lateral branch which
in turn leads directly to a basidium or to a new subhymenial cell; it follows
that the basidia -- arising as they do from extremely variable depths -- are of
very different lengths...." (translated by LLN). C. grossula also has a
candelabra-like subhymenium, but it is not as strongly developed. Josserand
(1955) drew attention to the similar hymenial development in C. chrysophylla
(as Omphalia) and C. grossula (as O. abiegna). He noted that this
developmental pattern led to an abundance of immature elements in the
hymenium, resulting in a low level of spore production. These observations
explain the difficulty in obtaining satisfactory spore deposits for either
species.
Oregon basidiomes of C. grossula produced scant spore deposits; a
thin deposit on paper is creamy-white; deposits on one microscope slide
appeared very slightly greenish. In his descriptions of the species (as O.
wynniae in 1882 and O. bibula in 1888) Quélet described a white or greenish
spore print ("hyaline ou verdatre"). After obtaining sufficiently heavy spore
deposits (only twice over a 21 year period), Josserand (1955) noted a very
pale ("trés, trés pale") creamy-greenish color when the print was compared to
a very white standard, bicarbonate of soda.
Possibly because of similarity in size, habit, overlapping habitats, and
color when faded, C. grossula has been confused with another clampless
species, Phytoconis ericetorum (Fr.) Redhead & Kuyper (also variously
known as Omphalina ericetorum (Fr.) M. Lange or Omphalia umbellifera
(Fr.) Kumm.). Older faded forms of both species can be microscopically
386
Figs. 3-5. Chrysomphalina grossula, Fig. 3. Basidiospores. (LLN 92.04.19-
1). Fig. 4. Candelabra-like subhymenium with irregularly shaped di-
/trichotomously branched hyphal elements giving rise to a dense hymenial
layer with basidioles and basidium. (LLN 93.04.18-1). Fig. 5. Femur-like
hyphal cells from pileal trama (LLN 93.04.18-1). Scale = 10 Um.
387
Figs. 6-7. Chrysomphalina grossula. Fig. 6. Exserted cystidioid end cells
from pileipellis (LLN 93.04.18-1). Fig. 7. Exserted cystidioid elements on
stipe (LLN 93.04.18-1). Scale = 10 um.
RRS
differentiated: hyphae in the pileal and lamellar trama of C. grossula are
markedly inflated (reaching 22 {1m in diameter) and constricted at the septa
(cf. Fig. 7) while those of P. ericetorum are narrower (4-10 tim) and not
normally swollen at the septa [cf. Bigelow, 1970]. Additionally, P. ericetorum
is lichenized with the basidiomes surrounded by a thallus composed of small
sphaerules. This species, unlike C. grossula, frequently develops a fairly
conspicuous pubescent vestiture on its stipe.
388
Choice of Species Epithet
The highly variable coloration and thickening lamellae of C. grossula
have contributed to the generation of several species epithets. Persoon (1828)
named the mushroom Agaricus grossulus, selecting a species epithet referring
to the gooseberry green coloration. The fungus was later described as a pale
yellow variety of Agaricus umbelliferus by Berkeley & Broome in 1875 (as
var. abiegnus), probably based on aged, partially faded specimens; the name
"abiegnus" refers to its normal substrate, coniferous wood. The species in its
lemon-yellow state, a coloration common in Hygrophorus sensu lato, was next
described as Hygrophorus wynniae by Berkeley & Broome in 1879 who
named it after the collector, Mrs. Lloyd Wynne. Apparently this assignment
to Hygrophorus obscured the fact that the authors had earlier described the
same agaric as an omphalioid taxon.
Quélet (1886) simultaneously assigned the species two names:
Omphalina umbellifera var. citrina for the more mature lemon-yellow colored
form, and Omphalina bibula, a reference to its bibulous nature, for the more
intense, younger olive-citrine colored form. In 1888 Quélet treated O.
wynniae as conspecific with O. bibula. Rolland (1891) named what he
believed to be a new yellow-colored form of the olive-citrine fungus
Omphalia bibula var. citricolor.
Lange (1930) transferred A. umbelliferus var. abiegnus to Omphalia
at the species level, treating H. wynniae, Omphalina bibula and O. umbellifera
var. citrina as synonyms. After examining the type of A. grossulus, Singer
(1961b) placed both O. abiegna and O. wynniae in synonymy with A.
grossulus, transferring the species to Omphalina. Clémencon (1982) accepted
Singer's synonymy but placed the taxon into Camarophyllus (q.v.). Both the
macroscopic description by Persoon (1828) and the microscopic description
of the type by Singer (1961b) are consistent with our concept of the species.
Agaricus chrysoleucus Pers.: Several authors have considered
Agaricus chrysoleucus Persoon (1801) to be an earlier name for the fungus
we call C. grossula. We do not accept this synonymy.
Lange (1930) treated A. chrysoleucus (as Omphalia umbellifera var.
chrysoleuca Pers.) as a synonym of O. abiegna, although he acknowledged
that there was confusion regarding Fries' concept of the latter. Konrad &
Maublanc (1934) listed Omphalia umbellifera var. chrysoleuca Pers. [non
Fries] as a synonym of Omphalia wynniae. They additionally recognized var.
bresadolae, listing O. bresadolae R. Maire, Clitocybe xanthophylla Bres., and
Omphalia hypoxantha Bres. as synonyms. However, A. chrysoleucus as
originally described by Persoon in 1801 has more characters in common with
the fungus currently named Gerronema strombodes in Europe than with C.
grossula. Persoon's A. chrysoleucus was said to be cespitose on conifer trunks
in the Harz Mountains. Colors of the pileus, lamellae, and stipe were
described as “albido... distantibus sulphureis...albo" respectively. Clitocybe
xanthophylla Bresadola (1881), considered synonymous with G. strombodes
(Beardslee 1917, Singer 1970, Clémencon 1982a, Breitenbach & Kranzlin
1991), was also described as cespitose on conifer trunks in the Italian Alps
389
(Bresadola 1881, 1928). Its protologue reads very much like that for A.
chrysoleucus: Pileus "griseocanus", lamellae "“distantes...laete flavae", and
stipe “albo-griseus". We believe A. chrysoleucus is more likely to be
synonymous with Clitocybe xanthophylla than with C. grossula. However,
even this synonymy is questionable (cf. below).
Fries (1818) appears to have misapplied the name A. chrysoleucus
Pers. to a species having broad yellow lamellae and occurring on bare earth in
Sweden, and subsequently (Fries 1821) applied the name A. chrysoleucus to
yet another species with a deeply umbilicate pileus and narrow lamellae. By
1830 Fries considered A. chrysoleucus a dubious species, possibly a form of
Agaricus melizeus. Later Fries (1832) listed A. chrysoleucus twice with two
different authorities, "Pers." and "Fr.", in the index to the Systema
Mycologicum, clearly indicating that he had earlier misapplied the name A.
chrysoleucus to a different fungus from that described by Persoon. In 1838
Fries no longer synonymized A. chrysoleucus Pers. with the hygrophoroid A.
melizeus, considering the former a yellow-gilled lignicolous form of A.
umbelliferus [possibly Phytoconis luteovitellina (Pilaét & Nannfeldt) Redhead
& Kuyper]; at this time he recognized A. chrysoleucus Fr. as a distinct taxon.
The lignicolous yellow-gilled form described by Fries (1838) may have been
either a Gerronema or Chrysomphalina.
Given this history of confusion, we prefer to drop the name A.
chrysoleucus from usage, considering it a nomen confusum.
Generic placement
The taxon under discussion has most often been treated as an
omphalinoid species. For instance, Bigelow included C. grossula in
Omphalina as O. wynniae. while Singer, after initially (1961b, 1962) placing
itin Omphalina as O. grossula, later incorporated the species in Gerronema
(Singer 1975, 1986). Bigelow and Singer's differing circumscriptions of
Omphalina and selected segregate omphalinoid genera will be fully explored
in) Party 2,
Some authors, however, have emphasized the similarities between C.
grossula and hygrophoraceous species. Clémencon (1982b) placed C.
grossula into Camarophyllus subgenus Aeruginospora because it appeared to
have a highly irregular hymenophoral trama and because the basidiomes
sometimes appear slightly greenish, linking it to the putatively green-spored
type of the subgenus. As typified by C. pratensis (Pers.: Fr.) Kumm.,
Camarophyllus is a typically hygrophoraceous genus with elongated basidia
characterized by having a distinctly irregular lamellar trama and a terrestrial
habit. Out of the five Camarophyllus sections accepted by Singer (1986),
only section Aeruginospora is characterized by clampless hyphae. Notably all
three species included by Singer in this section are not lignicolous, which
Clearly makes inclusion of the lignicolous C. grossula an anomaly.
Furthermore, C. grossula does not have a typically irregular lamellar trama. In
Our opinions, the species is not closely allied to C. pratensis and does not
belong in the Hygrophoraceae. Kuyper (1986) reached a similar conclusion
in his cladistic analysis of European omphalinoid genera.
390
CHRYSOMPHALINA CLEMENCON
Given the anatomical similarities between Chrysomphalina grossula
and C. chrysophylia outlined earlier, we consider the two species to be
congeneric. Chrysomphalina chrysophylla is the type species for the genus
Chrysomphalina Clg. (non Haas, see Redhead 1986). Clemencon (1982a)
validated Chrysomphalina to accommodate two omphalinoid species, C.
chrysophylla and C. strombodes (Berk. & Mont.) Cl¢. We interpret the genus
differently, restricting Chrysomphalina to species having a hymenium which
thickens over an extended life-span by the production of basidia of different
lengths on short subhymenial cells (see Clémengon 1982a, Kost 1986a,
Kiihner 1928). In C. chrysophylla the combination of hymenial thickening
and short subhymenial cells was so conspicuous that Clémencon coined the
term "pachypodial trama" for it. This tissue is formed, but to a lesser extent, in
C. grossula. A third species, C. aurantiaca (Peck) Redhead, also exhibits this
phenomenon; however, here the pachypodial trama is even less well-developed
than in C. grossula. We consider the development of this type of hymenium
to be not only taxonomically significant as emphasized by Clémencon, but
indicative of a possible linkage to the Cantharellaceae, as suggested by Kiihner
(1928) and Kost (1986b) (See also Arpin & Fiasson 1971 and Tyler 1971).
Chrysomphalina, as represented by the type species, also shares with
Cantharellus the presence of intracellular carotenoid pigments (Arpin &
Fiasson 1971, Kost 1986b, Kiihner 1980). Other important features of the
three Chrysomphalina species are monomitic tissues, strictly lignicolous habit
associated with white rot, and the absence of clamp connections on all tissues.
Included in the genus are five taxa -- the type species C. chrysophylla
with three varieties [C. chrysophylla var. chrysophylla, C. chrysophylla var.
salmonispora (Bigelow) comb. nov. (Basionym: Omphalina chrysophylla var.
salmonispora Bigelow. Mycologia 62: 22. 1970.), and C. chrysophylla var.
hoffmanii (Peck) stat. et comb. nov. (Basionym: Agaricus hoffmanii Peck, New York
State Mus. Rept. 24: 60. 1872.)], C. aurantiaca, and C. grossula.
Key to Chrysomphalina species
1. Basidiomes typically uniformly orange colored (pileipellis and
stipitipellis concolorous with context; only bright yellowish to rosy
orange pigments present); pileus margin fringed by hair-like
concolorous scales, at least when young.................... C. aurantiaca
1. Basidiomes typically bi-colored (pileipellis typically more darkly
colored than context; both an intracellular brownish or greenish
fuscous pigment and a yellowish or orange pigment present);
pileus either glabrous or subglabrous, or if scaly, with darker
scales confined,to the cemtral region |<. 0.5...,.cosssveorasevgenesusssses. vanes 2
2. Basidiomes bright greenish-yellow when young, becoming yellower
or paler with age but not exhibiting orangish hues at any
stage; tramal cells frequently femur-like at maturity (cf. Fig.
7); basidiospores with broadly rounded ends (cf. Fig. 3)
DAR TERS Ratt, MEET Nein NP my rer ak ot adieate C. grossula
2. Basidiomes with orange (orange, pinkish orange, or orangish
yellow), never greenish tints; tramal cells rarely femur-like;
basidiospores elongated, with many slightly tapered toward the
ADC XP CLOVE TOS) S7UA) etc were ste: Seve mere en nneteteres er tees coach cases 3
3. Pileipellis either glabrous or subglabrous, lacking dark scales on
CHS Cee Stace eitnns Mle okaer tence eterna C. chrysophylla var. hoffmanii
3. Pileipellis with definite dark scales on disc and inner margin................ 4
4. Basidiomes with orange or orangish yellow tints, lacking pinkish or
SALMON LONCS occ eats ek. C. chrysophylla var. chrysophylla
4. Basidiomes with pinkish-orange Or SalMOM tOMNES...........ceeeeeeeeeees
Relist ayes i ad A pe ng weed ae C. chrysophylla var. salmonispora
We believe C. chrysophylla var. salmonispora may be a color variant
with either different carotenoids or a different mixture of carotenoids from
those found in C. chrysophylla var. chrysophylla. Although C. chrysophylla
var. hoffmanii may be no more than a growth form, it is impossible to tell
from current data whether the smooth pilear condition is genetically or
environmentally controlled; some collections (e. g. Stuntz 9776) range from
the glabrous to the scaly condition. Additionally the spores are virtually
identical in all three varieties (Figs. 9-11) and differ subtly from C. grossula
(Fig. 3) and C. aurantiaca (Fig. 8).
A, YoQUBAN00|
Figs. 8-11. Basidiospores. Fig. 8. Cirysomphalina aurantaca(LLN 93.08.03-3).
Fig. 9. Chrysomphalina chrysophylla var. chrysophylla. (CArdrey 1429).
Fig. 10. C. chrysophylla var. hoffmaniit (SARedhead 7700). Fig. 11. C.
chrysophylla var. salmonispora (LLN 93.08.03-2b). Scale = 10 um.
392
Excluded species: In our classification, Chrysomphalina strombodes is
not congeneric with C. chrysophylla. Microscopical examinations of both
North American and European collections (see specimens examined) have
revealed that taxa identified as C. strombodes on both continents form
basidiomes composed of sarcodimitic tissues and therefore should be retained
in Gerronema. Additionally, they represent two taxa: a North American
species with minute dark brown to blackish appressed squamules on the pileus
composed of fascicles of clavate to clavate-pedicellate pileocystidia with
brown cellular contents (Fig. 14) and a European species with innately
radiating fibrils, wholly lacking pileocystidia and resulting squamules. As the
name Gerronema strombodes applies to the North American species (Fig. 12),
originally described from Ohio, another name must be used for the European
species (Fig. 13). Given our doubts about the name A. chrysoleucus, the next
available epithet is Clitocybe xanthophylla, for which authentic material has
been examined. We therefore propose for the European species the name
Gerronema xanthophyllum (Bres.) comb. nov. [(Basionym = Clitocybe
xanthophylla Bres. Fungi Tridenti Vol 1: 8. 1881. Synonyms = Omphalia hypoxantha Bres.
nom. nov. Icon. Mycol. VI. Tab CCLIX 1928. (2on Omphalia xanthophylla (Berk. & Curt.)
Sacc. Syll. Fung. 5: 312. 1887 = Agaricus (Omphalia) xanthophyllus Berk. & Curt. Ann. Mag.
Nat. Hist: 4. 1859.) = Omphalia bresadolae R. Maire nom. nov. Ann. Mycol. 11: 338. 1913.]
SPECIMENS EXAMINED Chrysomphalina aurantiaca: NORTH AMERICA:
CANADA: BRITISH COLUMBIA: Glacier Natl Park, Illecillewaet River --
IX.9.1980 Redhead 3533 (DAOM 184112, NEOTYPE). USA: CALIFORNIA:
Mendocino Co, Rockport -- X//.31.1968 JWLennox 150'b. WASHINGTON:
Chelan Co, Rainy Pass -- LLN 93.08.03-3, coll. JFAmmirati & TWKuyper; Clallam
Co, Olympic Natl. Pk., Sol Duc Falls -- LLN 92.10.14-4, LLN 92.10.14-7; King Co,
Redmond -- X/.24.1979, FVdBogart 3980. Lewis Co, Cispus -- X.22.1972 Stuntz
17500; San Juan Co, Friday Harbor -- X//.8.1974 SLibonati-Barnes 408; Yakima
Co, White Pass: Clear Lake -- V/.4.1989 JFA 9905 coll. LBaxter.
Chrysomphalina chrysophylla var. chrysophylla: NORTH AMERICA:
CANADA: ALBERTA: Kanaskis Valley, Marmot Creek Basin -- V//.13.1972
RMDanielson 308 (DAOM 143591). BRITISH COLUMBIA: Vancouver Island,
Pacific Rim Natl Park, Long Beach .-- [X.29.1979 SARedhead 3247 (DAOM
175260). QUEBEC: L'Islet co, Elgin Rd -- X.19.1949 HACJackson (DAOM 84909).
USA: OREGON: Coos Co, Beaver Hill Forest -- X.24.1986 CArdrey; [X.4.1989, C.
Ardrey 1429; Multnomah Co, Estacada -- LLN 91.12.30-4, coll. LLN & JRogers.
WASHINGTON: King Co, Seattle -- X/.12.1956 Stuntz 9776, coll. Bell & Stuntz;
Pierce Co, Mt. Rainier Natl Park, Lower Tahoma Creek -- /X.12.1948 Stuntz 4421, det.
AHSmith; San Juan Co, Friday Harbor -- [X.17.1979 FVdBogart 3920. WYOMING:
Teton Co, Flagstaff Road -- V//I.12.1987 JFAmmirati 9525.
Chrysomphalina chrysophylla var. hoffmanii: NORTH AMERICA:
CANADA: BRITISH COLUMBIA: Mt. Revelstoke Natl Park, Illecillewaet
Riv. -- [X.27.1980 SARedhead 4124 (DAOM 181086). USA: WASHINGTON: Ferry
Co, White Mountain, HWY 20 -- VII.1.1993, SAR 7700 coll. Ammirati, Norvell,
Redhead & Seidl.
Chrysomphalina chrysophylla var. salmonispora: NORTH AMERICA:
USA: WASHINGTON: Chelan Co, Rainy Pass -- LLN 93.08.03-2b, coll.
JFAmmirati.
Chrysomphalina grossula: See under species description above.
Bie 8)
Gerronema strombodes: NORTH AMERICA: USA: CONNECTICUT: New
London Co, Pachaug St. Forest. -- V/J/I.4.1988 coll. unknown [N.E. Mycol. Foray]
(DAOM 199314). NORTH CAROLINA: Swain Co, Great Smoky Mtns. Natl. Park --
VII.19.1988 D.E. Desjardin 4616 (DAOM 199312).
Gerronema xanthophyllum: EUROPE: ITALY: --190] JBresadola (ex herb
Bresadola, NY). SWITZERLAND: Lucerne Cn: Ruswil, Rotbachtobel -- V/. 15.198]
JBachler 1506-81 BA 5 (LU); Adlingenswil, Meggerwald -- V//.13.1989 FKranzlin 1307-
89K 1 (LU).
14
Fig. 12. Gerronema strombodes. Basidiospores. (DAOM 199314), Fig. 13.
Gerronema xanthophyllum Basidiospores. (LU FK 1307-89 K1). Fig. 14.
Pileocystidia in Gerronema strombodes. A. (DAOM 199314). B. (DAOM
179312), Scale = 10 um.
394
PART 2: OMPHALINA sensu BIGELOW
Omphalina was erected by Quélet (1886) for white-spored, centrally
Stipitate mushrooms with thin umbilicate or tubaeform pilei, decurrent
lamellae, and cartilaginous stipes. From this macroscopical circumscription
the terms "omphalioid" and "omphalinoid" were derived, i.e. "any mushroom
[of small stature ~ 1-2 cm diam.] with decurrent or subdecurrent gills, a
cartilaginous stipe, a broadly convex to depressed pileus, a slightly to
markedly depressed pileal disc, and lacking an annulus and a volva" (Largent
& Baroni 1988),
Microscopical observations, however, suggest that this is a polyphyletic
group, with parallel evolution having produced morphologically similar but
anatomically distinct taxa. This view is consistent with recent segregations of
genera from Omphalina sensu lato by taxonomists apparently seeking
monophyletic genera within a more natural classification system.
Chrysomphalina, one such segregate genus, has already been outlined
above. Other recently described or redefined genera (i.e. Gerronema,
Pseudoarmillariella, Haasiella, Rickenella, and Phytoconis) are discussed
below, followed by an evaluation of the genus Omphalina itself. A key to the
genera as we accept them and expanded generic descriptions follow this
discussion.
GERRONEMA Singer: Singer (1951b) erected the genus Gerronema
to accommodate three tenacious omphalinoid to clitocybeoid species from
South America. He later (1964) transferred some species traditionally placed
in Omphalina to Gerronema. Problems have arisen from the different
circumscriptions of Omphalina by Singer and Bigelow. Virtually all the
species recognized in Bigelow's 1970 monograph of Omphalina are included
in Singer's expanded concept of Gerronema (Singer 1986), while Bigelow's
concept of Clitocybe (Bigelow 1982a&b, 1985) encompassed any species left
by Singer (1986) in Omphalina. In both classification systems, the primary
distinction between the paired genera, either Omphalina sensu Singer and
Gerronema sensu Singer or Clitocybe sensu Bigelow and Omphalina sensu
Bigelow is pigment-based. Singer distinguishes Omphalina from Gerronema
almost solely on the presence of fuscous, intraparietal or incrusting pigments,
present in the former and absent in the latter. Bigelow’s circumscripton of
Omphalina revolves around the presence or suspected presence of carotenoids
in all included species. Neither author analyzed pigments in the species each
treated.
Pigmentation evidently either carried more weight than other
taxonomic characters or appeared to be the strongest taxonomically important
feature in the absence of other observed characters. However, Gerronema
sensu Singer (1961la, 1964, 1975, 1986) is considered by many authors to be
heterogeneous (i.e. Clémengon 1982, Moser 1983, Kuyper 1986), and several
attempts have been made to reduce it to a more natural group. We agree
Gerronema sensu Singer is heterogeneous and believe that Omphalina sensu
Singer (1986) is also heterogeneous, but to a lesser extent. Lange's (1981.
See also 1992) expansion of Omphalina to include Gerronema, Haasiella,
395
Rickenella, and Phaeotellus, thus reflecting the historical basis of Omphalina,
is appealing because of its simplicity, but clearly, it also represents a
polyphyletic grouping.
In the case of Omphalina sensu Singer (1986), emphasizing
pigmentation based primarily upon a generalized pathway leading to
melanization (see Ellis & Griffith 1974; Hegnauer et al. 1985; Swan 1974)
appears to us to result in an artificial classification. Notably, there are many
genera in the Agaricales sensu Singer and in the Cantharellaceae that include
species with more than one pigment group. In our opinion, dependence upon
the presence or absence of especially common pigment groups, such as
melanin, to define genera is perhaps unwise; in this case, pigmentation should
be used judiciously or in conjunction with other taxonomic features. We
believe that a combination of anatomical and biological characters clearly
requiring a series of complex genetic expressions are of greater importance.
Molecular analyses of DNA, such as that being conducted by Lutzoni and
Vilgalys (1993), would be even more instructive.
The fact that the basic structural tissue forming the basidiomes of
certain genera is sarcodimitic -- in contrast to monomitic, dimitic, or trimitic --
is of fundamental taxonomic importance, because the formation of
sarcodimitic tissues is a complex series of processes requiring the expression
of many genes. However, the extent to which sarcodimitic tissue formation
can be used as an important taxonomic indicator of different evolutionary
pathways lies somewhere between the extremes at the generic (i. e. 7rogia, cf.
Corner 1966, 1991) or family (i.e. Xerulaceae, cf. Redhead 1987) levels and
schemes in which the presence of
sarcodimitic tissue is not taken into
account, ignored, or dismissed (Singer
1986, Reijnders 1993). Sarcodimitic
tissue formation can be used to help
define genera or generic groupings
(Vilgalys & Rehner 1993).
a Redhead (1986) restricted the genus
Gerronema to species having
sarcodimitic tissues. Subsequent
ie xcontizination= Ol vesarcodimitic
construction in G. melanomphax Sing.
(the type of the genus) was made by
Redhead from an isotype (Singer T-
1094 at MICH; Fig. 15). Species
lacking sarcodimitic tissues and
transferred to Gerronema after its
inception are thereby excluded. By
further delimiting the genus in this
way, a more natural group is defined.
Fig. 15. Gerronema melanomphax (Isotype Singer T-1094 (MICH).
Basidium, basidiole, and, basidiospores. Scale = 10 um.
396
PSEUDOARMILLARIELLA (Singer) Singer: Bigelow (1982a)
transferred Agaricus ectypoides Peck, the type for Pseudoarmillariella, to the
genus Omphalina. By including A. ectypoides in Omphalina, he radically
modified his 1970 concept of the genus which excluded species with amyloid
spores. Although he did not so specify in published articles, Bigelow stated to
Redhead in 1985 that he based this transfer upon the many similarities
between P. ectypoides and C. chrysophylla.
The monotypic genus Pseudoarmillariella is characterized by the
production of a thickened hymenium and subhymenium (Singer 1956). Its
subhymenium is said to be "subirregularly intermixed-subramose, its elements
short, strongly interlaced-curved in all directions and therefore at times
appearing cellular (much like the subhymenium of Cantharellula)" (Singer
1986). Pseudoarmillariella ectypoides differs from all Chrysomphalina
species by the production of incrusting pileal pigments, amyloid spores and
the formation of clamp connections. This indicates a sufficient hiatus to
recognize Pseudoarmillariella as distinct from Chrysomphalina.
HAASIELLA Kotlaba & Pouzar: Haasiella venustissima (Fr.) Kotl.
& Pouzar (1966), the type of the genus, and Chrysomphalina chrysophylla
were originally considered to be congeneric by Haas who invalidly proposed
the genus “Chrysomphalina Haas" in 1962. Clemengon (1982a) excluded H.
venustissima from Chrysomphalina Clg. when he validated the genus.
Like Chrysomphalina and Cantharellus, Haasiella possesses
carotenoid pigments (Arpin & Fiasson 1971). However, it differs from
Chrysomphalina by the formation of thick, metachromatic basidiospore walls
(Kotlaba & Pouzar 1966) and a slightly gelatinized pileipellis (Kost 1986b).
Both recognized species of Haasiella are distinctly terrestrial in contrast to the
lignicolous Chrysomphalina species, and one, H. splendidissima Kotl. &
Pouz., forms clamp connections.
Singer (1986) recognized Haasiella, but only as a section of
Gerronema. However, since basidiomes of both species in his section are
composed of monomitic tissues, the species are excluded from Gerronema in
our classification. Haasiella is only known from Europe and was not treated
by Bigelow (1970) in his North American monograph. Lange (1981)
includes this genus in Omphalina sensu lato.
RICKENELLA Raithelhiiber: Although not accepted in Omphalina
by Bigelow, he questioned generic placement of the type species, R. fibula
(Bull. : Fr.) Raithelh. (Bigelow 1970). Rickenella was distinguished from
Gerronema sensu lato by Raithelhiiber (1973) based on the presence of
prominent characteristically shaped leptocystidia abundantly covering the
pileus, stipe, and hymenial surface. These same features serve to distinguish
Rickenella from Omphalina. Rickenella has been widely accepted by
European and North American agaricologists (notably Kost 1984, Lamoure
1979) and we accept the genus as presently circumscribed.
All Rickenella species are small, omphalinoid, and bryophilous; the
species are known to colonize either mosses or liverworts (Redhead 1981, Kost
397
1984) and pure isolates form distinctive slow-growing colonies (Lamoure
1979). This complex of features amply characterizes the genus. Additionally
Kost (1984) has drawn attention to the physalomitic nature of the tramal
tissues in this genus (physalomitic hyphae = conspicuously inflated hyphae
deeply indented at the septa, see Clémencon 1982a).
Taxa now assigned to Rickenella have been placed into various genera
by different authors attempting to create a natural classification. The two
most common species, R. fibula (Bull.: Fr.) Raithelh. and R. swartzii (Fr.)
Kuyp., were included in Omphalina at its inception (Quélet 1886), and Lange
again accepted this generic placement in 1981. However, Kiihner (1938),
Smith (1947), and Bigelow (1970) included Rickenella species in the genus
Mycena. In 1943 Singer treated them in his genus Hemimycena, but by 1951
he considered Hemimycena congeneric with Marasmiellus, including its
cystidiate species in Marasmiellus, sect. Fibulae. Subsequently Singer (1961)
transferred the species in section Fibulae to the genus Gerronema. When
Singer (1962) resurrected Hemimycena as a distinct genus, he left section
Fibulae in Gerronema. We recognize this section as the genus Rickenella.
PHYTOCONIS Bory de St. Vincent: As circumscribed by Redhead
& Kuyper (1987, 1988), Phytoconis is comprised of species previously treated
in Gerronema and Omphalina (Singer 1986). Phytoconis was delimited
primarily on the basis of lichenization and secondarily by the presence of a
leptomitic hyphal system (used by Kuyper, 1986). Molecular analysis of the
25S rRNA gene of these lichenized species indicates that Phytoconis is a
monophyletic genus linked to Omphalina (Lutzoni & Vilgalys 1993).
Notably, the genus includes both species with intracellular bright
pigments and species with intraparietal and incrusting melanized pigments.
The complex, unique, and characteristic nature of the fungal sheaths formed
by these species (as detailed by Redhead & Kuyper, 1987) offers ample
evidence of the close relationship among them. Additionally the
subhymenium in Phytoconis species is loosely structured due to the
elongated subhymenial cells and lack of clamp connections. In this regard
Phytoconis differs from Omphalina sensu Stricto.
OMPHALINA Quélet: Although this generic name is central to the
discussion of omphalinoid genera, its usage is the most controversial and
unstable. A major problem lies in the typification of the name. The first
lectotype was Omphalina hydrogramma (Fr.) Quél., selected by Earle (1909);
however, this typification could be superseded by O. umbellifera (Fr.: Fr.)
Quél. (proposed by Singer & Smith, 1946; Jgrgensen & Ryman, 1989), by O.
epichysium (Pers.: Fr.) Quél. (proposed by Redhead & Weresub, 1978;
Redhead & Kuyper, 1993), or by O. pyxidata (Pers.: Fr.) Quél (submitted as
an alternative by Redhead, 1993).
We have provisionally adopted O. pyxidata as type of the genus since
it has to date received the strongest support in the Committee for Fungi and
Lichens of the International Association for Plant Taxonomists. We restrict
Omphalina to a primarily bryophilous group of species centered around O.
pyxidata.
398
Omphalina species generally lack distinctive hymenial cystidia and
highly differentiated cuticles (but exceptionally can have cystidioid end cells
or scales, i.e. O. sphagnicola (Berk.) Moser). Omphalinas have smooth, thin-
walled, inamyloid spores. Anatomically Omphalina differs from both
Chrysomphalina and Phytoconis by the relatively narrow, compact
subhymenium composed of short basidia-bearing cells; this tissue, which
binds the hymenium more tightly than does the subhymenium of Phytoconis,
is best observed in squash mounts. Therefore, basidia and basidioles in
Phytoconis species tend to spread out relatively easily while those of
Omphalina tend to spread out in rafts or sections of hymenium. We include in
Omphalina several species groups largely separated by differences in
pigmentation. We do not restrict the genus to species with darkly melanized
incrustations or intraparietal pigments as does Singer (1986). Examples of
differently pigmented species are: O. postu (Fr.) Singer (orangish intraparietal
pigments), O. viridis (Hornem.) Kuyper (blue-green/blackish incrusting
pigments), O. pyxidata (brownish incrusting pigment) and Omphalina
hohensis (Smith) comb. nov.* (greyish black incrusting pigments) *Basionym
= Omphalia hohensis A.H. Smith, Contr. Univ. Mich. Herb. 5: 27. 1941. Delimitation
from Arrhenia Fries (including Phaeotellus Kihner & Lamoure and
Leptoglossum Karst. cf. Redhead 1984) is open to reevaluation using
molecular analysis.
DISPOSITION OF OMPHALINA SENSU BIGELOW 1970
Most of the twelve species and one variety Bigelow (1970) included in
Omphalina are placed elsewhere following the generic classification for
omphalinoid agarics outlined above.
As discussed in Part 1, five taxa belong in Chrysomphalina: O.
chrysophylla (Fr.) Murrill var. chrysophylla = C. chrysophylla var.
chrysophylla, O. chrysophylla var. salmonispora = C. chrysophylla var.
salmonispora, O. hoffmanii = C. chrysophylla var. hoffmanii, O. luteicolor
Murrill = C. aurantiaca (Redhead 1986), and O. wynniae = C. grossula.
Four other species were transferred to Phytoconis by Redhead &
Kuyper (1987, 1988): O. ericetorum = P. ericetorum, O. luteovitellina (Pilat.
& Nannf.) M.Lange = P. luteovitellina, O. hudsoniana (Jenn.) Bigelow = P.
viridis, and O. sphagnophila (Peck) Bigelow = P. ericetorum.
Omphalina subclavata (Peck) Murr. is now Gerronema subclavatum
(Peck) Redhead (Redhead 1986).
Of the remaining three species treated by Bigelow, Omphalina postii
(Fr.) Sing. remains in the genus Omphalina. We do not consider the tissues in
the stipes of O. postii or the related species Omphalina marchantiae(Sing. &
Cl¢.) comb. nov. (Basionym = Gerronema marchantiae Sing. & Clg. Schweiz.
Zeitschr. Pilzk. 49: 118. 1971.) to be sarcodimitic as suggested by Senn-Irlet et
al. (1990). Omphalina marchantiae has been reported from Alaska (Laursen
& Ammirati 1982, Laursen & Chmielewski 1982) and British Columbia
(Kroeger 1989) in North America since Bigelow's 1970 monograph.
399
Two species of Omphalina sensu Bigelow -- O. occidentalis (A.H.Sm.)
Big. and O. olivaria (Peck) Sing. -- remain, both of uncertain status.
When Smith (1941) first described Omphalia occidentalis, he
compared it with both Mycena and Hygrophorus, noting its ambiguous
taxonomic affinities. Bigelow (1970) also felt that O. occidentalis was poorly
delimited from Hygrocybe and arbritrarily retained it in Omphalina. It
appears to us that this species may be closely allied to species such as
Hygrocybe parvula (Peck) Murr. and Hygrophorus (section Hygrocybe)
mycenoides A.H.Sm. & Hesl., both of which have small yellow basidiomes
with length:width basidia ratios that overlap those for O. occidentalis (cf.
Hesler & Smith 1963). The presence of clamp connections and a thin ixocutis
in O. occidentalis, both present in H. parvula and H. mycenoides, are features
more consistent with Hygrocybe than with the omphalinoid genera. Bas
(1988) noted that strict application of length-width ratios does not apply for
many Hygrocybe species, and reliance must be made on other features such as
bright colors and viscid pileal surfaces, features particularly evident in O.
occidentalis. Thus, we propose the name Hygrocybe luteo-omphaloides nom.
nov.* for this fungus. *Basionym = Omphalia occidentalis A.H. Smith, Contr. Univ.
Mich. Herb. 5: 28. 1941 nec Hygrocybe occidentalis (Dennis) Pegler in Pegler & Fiard, Kew
Bull. 32(2):310. 1978. nec Hygrophorus occidentalis Sm. & Hesl. Lloydia 2:18. 1939
The type of Omphalina olivaria possesses incrusting pigments in the
pileipellis and may in fact be a small Clitocybe as suggested by Bigelow
(1970). It is known only from the type collected over 100 years ago from a
burn in New York State. The name is retained in Omphalina on the
assumption it was associated with pyrophilous mosses. Bigelow (1970) has
already recorded the fact that other reports of O. olivaria are based on
misdeterminations.
KEY TO SELECTED OMPHALINOID GENERA
1. Cystidia present, densely and evenly distributed over pileipellis, hymenial
surfaces, and stipitipellis, visible under a hand lens as small hyaline hairs,
usually capitate, not fasciculate [bryophilous, lacking incrusting pigments,
SHORES INGIMNY| OLA CANG MNIN-WAIEDY ice cicccrnes ss ccses ves op dace oases senndes Rickenella
1. Cystidia lacking in most species; if present, more limited in distribution,
often inconspicuous or poorly differentiated, not capitate, sometimes
fasciculate in small squamules [other features variable] .........ccccccccceccceees 2
2. Spores amyloid [both incrusting pigments and non-incrusting pigments
present simultaneously, lamellar trama complex with thick dense lateral
subhymenial zones, hymenium thickening and basidia of different lengths,
LEONI CONUS CMa N. 1 < teal lan hare Wty UU OM Saeko | he Pseudoarmillariella
Pa Sporesmuamyloid, oer featuresavartaDle ein ne vadeav is Oks ent es veranda 3
400
3. Context of pileus and/or stipe composed of typical sarcodimitic tissues
[spores thin-walled, lignicolous, (primarily found in sub-tropical/tropical
FOQLONS) Jas ic taeroactetee tl cane see ena (oH anny bs oa dns Mecntoe te Batwa tet Gerronema
3. Context of neither stipe nor pileus composed of sarcodimitic tissues [other
features variable} 0 LAO ae, ROM ee eee 4
4. Spore walls conspicuously thickened and metachromatic in cresyl blue
[basidiomes orange, possessing carotenoids, terrestrial.
EXTRALIMITAD) <2 European} acne A ete Haasiella
4. Spore walls thin, not metachromatic [other features variable].............. 5
5. Thalli lichenized, composed of minute sphaerules or squamules while
superficially appearing to be terrestrial, bryophilous or lignicolous; walls
of hyphae which connect sphaerules or irregular granules and basal
mycelium noticeably thickened from slight to > 0.5 um [clamp
connections lacking, context of basidiomes leptomitic, subhymenial layer
a loose structure not forming a sealed compact membrane) ...... Phytoconis
5. Thalli not lichenized, typically mycelial and mostly embedded in substrate
(terrestrial, bryophilous, lignicolous); walls of basal mycelium not
thickened /[other featiires variable] 4% vita ees. siemens ay eee 6
6. Basidia of +/- conspicuously unequal lengths with the subhymenium
+/- developing into a pachypodium, hymenium thickening with age
[basidiomes brightly coloured by carotenoids {yellow, orange, yellow
mixed with green}, lignicolous, clamps lacking] .......... Chrysomphalina
6. Basidia more or less terminating at the same level, subhymenium a thin
layer; hymenium not thickening in age [other features
variable Faris 8 roto cee es Made oe ey ee 7
7. Basidiomes brightly pigmented and pileipellis gelatinized to subgelatinized
with a thin but distinct cutis [pileus minute, clamp connections present,
DryOpHLOUS eed he tara ee Hygrocybe luteo-omphaloides
7. Basidiomes not both brightly pigmented and with a gelatinized pileipellis
[other features Variable]... ic tciiiss cies tented ete det nes ee ne 8
8. Bryophilous or phycophilous (sometimes appearing terrestrial or
lignicolous due to sparse growth of associated algae or bryophytes),
never on sound clean wood or conspicuously causing white or brown
wood rots [with incrusting or intraparietal pigments, odor never
farinaceous, subhymenium a compact dense thin layer totally sealing
in the lamellar trama from the hymenium]............0.0.cc006 Omphalina
401
{A completely pigmentless species associated with algal films on silty
soil is Hemimycena ignobilis (Josserand) Singer, of debatable generic
disposition}
8. Terrestrial or lignicolous, not dependent on living algae or bryophytes;
if on wood mycelium clearly penetrating and decaying wood [easily
cultured on standard general media from spores, odor sometimes
farinaceous, many species large and not omphalinoid)].......... Clitocybe
GENERIC CIRCUMSCRIPTIONS
CHRYSOMPHALINA Clc.
Basidiomes fleshy, pileus convex, usually umbilicate; lamellae
decurrent, occasionally forking; stipe central and cartilaginous to fleshy;
spores white to slightly pigmented in mass, lacking a germ pore, walls thin,
smooth, inamyloid, hyaline, and not metachromatic in Cresyl Blue.
Pileipellis relatively undifferentiated from the monomitic tramal
tissues, often with projecting loose undifferentiated hyphal ends +/- aggregated
into small scales; often containing intracellular darkish pigments in addition to
cytoplasmic carotenoids in the tissues; tramal hyphae thin- to somewhat thick-
walled, inamyloid, nongelatinized, without clamp connections; hymenophoral
trama bidirectional; hymenium thickening, basidia in older specimens
conspicuously varying in length, subhymenium a thick layer forming a dense
band of tissue in age; lacking hymenial cystidia; sometimes with scattered
relatively undifferentiated cystidioid elements on the stipe
Lignicolous.
Type species: C. chrysophylla (Fr.) Cl¢
GERRONEMA Sing.
Basidiomes membranous to elastic, pileus convex to infundibuliform
or umbilicate; lamellae decurrent; stipe central, elastic to fleshy or
cartilaginous; spores white in mass, lacking a germ pore, walls thin, smooth,
inamyloid, hyaline, and not metachromatic in Cresy] Blue.
Pileipellis poorly differentiated, monomitic, often with intracellular
pigments differing from those in the trama or hymenium, occasionally with
scattered cystidioid elements on the disc; tramal tissue sarcodimitic (hyphae
basically of two types: elongated fusoid cells bonded by filamentous branched
hyphae, most easily observed in the stipe), inamyloid, +/- gelatinized, and
lacking dark intraparietal or incrusting pigments, with or without clamp
connections; hymenophoral trama sarcodimitic; hymenium not conspicuously
thickening.
Lignicolous.
Type species: G. melanomphax Sing.
402
HAASIELLA Kotl. & Pouz.
Basidiomes fleshy, pileus convex, slightly umbilicate; lamellae
decurrent, cantharelloid; stipe central and somewhat cartilaginous; spores
yellow to salmon rose in mass, lacking a germ pore, walls thickened (double
walls present), smooth, inamyloid, hyaline, red metachromatic in Cresyl Blue.
Pileipellis a layer of gelatinized, leptomitic hyphae often with swollen
hyphal ends; containing intracellular carotenoid pigments in the tissues; tramal
hyphae thin- to somewhat thick-walled, inamyloid, +/- gelatinized, with or
without clamp connections; hymenophoral trama irregular, possibly
bidirectional; subhymenium continually producing basidia; hymenium
thickening; lacking hymenial cystidia.
Terrestrial.
Type species: H. venustissima (Fr.) Kotl. & Pouz.
OMPHALINA Quél.
Basidiomes membranous to fleshy, pileus convex to infundibuliform
or umbilicate; lamellae decurrent, regular to cantharelloid; stipe central to
eccentric but not lateral, cartilaginous, most often glabrous; spores white in
mass, lacking a germ pore, walls thin, smooth, inamyloid, hyaline, and not
metachromatic in Cresyl Blue.
Pileipellis relatively undifferentiated from the monomitic tramal tissue
in most species, sometimes with aggregated hyphal ends forming small scales
usually restricted to the pileal disc, rarely abundant enough to form an
interrupted turf of fasciculate cystidioid elements; often pigmented with dark
or brightly colored intraparietal, incrusting, or intracellular pigments; tramal
hyphae thin-walled, inamyloid, nongelatinized, usually with but some species
without clamp connections; hymenophoral trama bidirectional to irregular;
basidia not varying in length; hymenium not appreciably thickening;
subhymenium a relatively thin, compact dense layer; poorly differentiated
hymenial cystidia present in at least one species concentrated near the lamellar
edge; caulocystidia, when present, generally poorly differentiated and ranging
from sparse to aggregated in small squamules.
Either bryophilous or lignicolous.
Type species: O. pyxidata (Pers: Fr.) Quél. subject to conservation.
PHYTOCONIS Bory
Basidiomes fleshy to membranous, pileus convex, usually umbilicate;
lamellae decurrent; stipe central, cartilaginous to tough, usually pubescent,
spores white in mass, lacking germ pore, walls thin, smooth, inamyloid,
hyaline, and not metachromatic in Cresyl Blue.
Pileipellis relatively undifferentiated from the leptomitic tramal tissue;
hyphae thin to somewhat thickened, inamyloid, nongelatinized, with either
fuscous intraparietal pigments or brightly colored intrahyphal pigments,
without clamp connections; hymenophoral trama sub-irregular; basidia in
older specimens varying in length; hymenium usually slightly thickening with
a loose subhymenial structure; lacking cystidia on the pileus and in the
hymenium; frequently with cystidioid or hair-like hyphal ends on the stipe.
403
Lichenized, characterized by thalli totally enveloping Coccomyxa algal
cells in non-perforated sheaths of polygon-shaped cells, forming either
scattered sphaerules or irregular granules typically less than 1 mm in diameter
or flattened/lobed thalli typically less than 1 cm in diameter connected by
filamentous hyphae.
On soils, mosses, peat, or decayed conifer wood.
Type species: P. botryoides (L.) Bory = P. ericetorum (Fr.) Redhead
& Kuyper.
PSEUDOARMILLARIELLA Sing.
Basidiomes fleshy, pileus convex, usually umbilicate; lamellae
decurrent, often forked; stipe central and cartilaginous to fleshy; spores white
to slightly pigmented in mass, lacking a germ pore, walls thin, smooth,
amyloid, hyaline, and not metachromatic in Cresyl Blue.
Pileipellis relatively undifferentiated from the monomitic tramal
tissues; with incrusting pigments and non-incrusting cytoplasmic pigments
simultaneously present; tramal hyphae inamyloid, +/- gelatinized, thin- to
somewhat thick-walled, with clamp connections; hymenophoral trama
subirregular; hymenium thickening, basidia in older specimens conspicuously
varying in length, subhymenium a dense layer with short, intermixed
subramose, strongly interlaced elements, curved in all directions; lacking
hymenial cystidia; stipe densely covered with thin-walled hair-like hyphal
ends.
Lignicolous.
Type species: P. ectypoides (Peck) Sing.
RICKENELLA Raithelh.
Basidiomes fleshy, pileus convex, usually umbilicate; lamellae
decurrent; stipe central and cartilaginous; spores white in mass, lacking a
germ pore, walls thin, smooth, inamyloid, hyaline, and not metachromatic in
Cresyl Blue. |
Pileipellis relatively undifferentiated from the physalomitic tramal
tissue; hyphae thin-walled, inamyloid, nongelatinized, and lacking dark intra-
parietal or incrusting pigments, with or without clamp connections;
hymenophoral trama regular; hymenium not thickening; characterized by
abundant capitate to non-capitate refractive-walled evenly scattered
leptocystidia on the pileus, hymenial surfaces, and stipe.
Bryophilous.
Type species: R. fibula (Bull.: Fr.) Raithelh.
Acknowledgments: We thank Drs. D. Lamoure (Lyon, France), R. Singer (F, Chicago)
and R. Watling (E, Edinburgh) for valuable notes and observations and Drs. Th. W. Kuyper
(Netherlands), J. Ginns (DAOM, Ottawa), and Mr. G. T. Norvell for critically reviewing the
manuscript. We also thank the curators of the following herbaria for the generous loan of
specimens: Dr. R. Watling (E), Dr. G. W. Mueller (F), Dr. D. L. Largent (HSC), Drs. J.
Breitenbach & F. Kranzlin (LU), and Drs. R. Fogel and R. Shaffer (MICH). Support to the
senior author by the Daniel E. Stuntz Memorial Foundation was greatly appreciated. Use of the
Cascade Head Forest Research Station, OR, as field support was provided by Dr. Sarah E.
Greene. Special thanks go to Mr. G. T. Norvell and Mr. O. L. Norvell for innumerable little
yellow mushroom sightings.
404
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---------- . 1961b. Type studies on basidiomycetes. X. Persoonia 2(1): 29-30.
~--------- . 1962. The Agaricales in Modern Taxonomy. ed. 2. Weinheim: J. Cramer.
a--------- . 1964. Die Gattung Gerronema. Nova Hedwigia 7: 53-92.
---------- . 1970. Flora Neotropica: Monograph No. 3 -- Omphalinae New York:
Hafner Publishing Company. 3-84.
wo nnen---- . 1975. The Agaricales in Modern Taxonomy, ed. 3. Vaduz: J. Cramer.
an-------- . 1986. The Agaricales in Modern Taxonomy, ed. 4. Koenigstein/Taunus:
Koeltz Scientific Books.
407
Singer, R. & A.H. Smith. 1946. Proposals concerning the nomenclature of the gill fungi
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240-299.
Smith, A.H. 1937. Notes on agarics from the western United States. Bull. Torrey Bot.
Club 64: 481
w--+------ . 1941. Studies of North American Agarics -- I, Contr. Univ. Mich. Herb. 5: 28.
---------- . 1947. North American species of Mycena. Ann Arbor: University of
Michigan Press. 1-521.
Swan, G. A. 1974. Structure, Chemistry, and Biosynthesis of the Melanins. /n Fortschritte
der Chemie organischer Naturstoffe, Herz, Grisebach & Kirby, eds. New
York: Springer-Verlag. 522-582.
Tyler, V.E. 1971. Chemotaxonomy in the Basidiomycetes. In Evolution of the Higher
Basidiomycetes, R.H. Petersen, ed.; Knoxville, U of Tennessee Press. 29-62.
Vilgalys, R. & S.A. Rehner. 1993. A preliminary phylogenetic analysis of several genera in
the Tricholomataceae based on nuclear large subunit ribosomal RNA sequence data.
Inoculum: Newsl. of the Mycol. Soc. Am44(2): 62.
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Volume L, pp. 409-427 January-March 1994
ALTERNARIA THEMES AND VARIATIONS (106-111)
EMORY G. SIMMONS
717 Thornwood Road, Crawfordsville, IN 47933
ABSTRACT
This paper reviews Alternaria infectoria and compares
with it A. triticimaculans Simmons & Perell6, sp. nov. (on
Triticum aestivum, Argentina), A. oregonensis Simmons,sp.
nov. (on T. aestivum, Oregon, U.S.A.), and A. metachromatica
Simmons, sp. nov. (on T. aestivum, Australia), all members
Otlecie. Av imrectoria ‘species group, \ Avo triticina and ‘A.
triticicola, both from T. aestivum, India, are reviewed for
contrast and clarification of taxonomic relationships in
Alternaria.
XIV. SOME ALTERNARIA TAXA ON TRITICUM
Introduction
The receipt of Alternaria specimens related to a leaf-
spot disease of wheat currently under study in Argentina and
a need for identification of the pathogen have required a
return to consideration of taxa similar to A. infectoria
Simmons (Simmons 1986). The phrase "Alternaria infectoria
complex" is used here as a grouping term for relatively
small-spored Alternaria taxa that are distinguished as much
by the prominence of their secondary conidiophore structure
as they are by the morphology of their conidia.
In the absence of their ascosporic teleomorphs, the
Alternaria taxa of this species complex are recognized in
culture primarily as loose to dense clumps of geniculate and
branching primary and secondary conidiophores with spore
chains whose conidial elements are separated by sturdy, often
elongate, readily distinguishable pseudorostrate connectives.
The nature of this sporulation pattern recently has been dis-
410
cussed and contrasted in detail with other kinds of Alterna-
ria sporulation patterns (Simmons and Roberts 1993).
The A. infectoria complex currently comprises three
named Alternaria species, all known as isolates derived
directly from ascospores of their teleomorphs: A. conjuncta
Simmons known at present from Pastinaca and Plantago; A.
infectoria, from Elymus, Festuca, Lolium and Triticum; and
A. ethzedia Simmons, from Brassica. The distinctive sporula-
tion pattern, which is determined by the strongly pseudoros-
trate nature of many of the conidia, is seen very frequently
among isolates of relatively small-spored taxa from numerous
substrates, sometimes as infecting agents but also probably
as saprophytes, and apparently without climate zone restric-
tion. All named species known until now for the complex are
anamorphs of Lewia Barr & Simmons (Simmons 1986).
Sorting out this entire group on the basis of cultures
is not high on my priority list, for members of the A. infec-
toria complex, even when culturally quite stable and segre-
gatable into recognizable taxa when studied side-by-side, are
highly resistant to verbal disaggregation. An additional
deterent resides in the fact that field material rarely
accompanies derivative cultures, so it is nearly impossible
to link modern isolates with any of the multitude of field-
based, potentially usable types and names in the literature.
It presumably will require an epiphytotic or industrial need
of significant proportions to divert molecular study to such
classification problems. However, I think that such an
approach eventually will be needed in sorting out the taxa
of this and other complexes of small-spored Alternaria
species.
The isolates from wheat in Argentina, mentioned above,
are supported by excellent sporulating field material. They
need identification; isolates of a similar taxon from wheat
in Oregon deserve attention; and another similar isolate,
from wheat in Australia, unique in its blue conidia that turn
pink in lactic acid, invites comparison and comment. All
three of these taxa have characteristics of the A. infectoria
species complex and are discussed here in comparison with
typical A. infectoria. Two other very distinctive Alternaria
species, A. triticina Prasada & Prabhu (1963) and A. triti-
cicola V. G. Rao (1964), originally described from wheat but
considered not to be morphological members of the A. infecto-
ria complex, are included here for purposes of contrast and
of guidance in identification of these Triticum inhabitants.
411
Conventions used in this discussion include: herbarium
and culture collection abbreviations are from Holmgren et al.
(1990), given here in boldface type; references to my records
of material studied are in record volume-page format, e.g.,
EGS 27-193; a contraction "Alt.T&V" is used to simplify ref-
erence to this series of papers on Alternaria, usually with
a number specifying a pertinent unit in the series (Simmons
1981-1993, all in Mycotaxon volumes of the period); routine
culture conditions include PCA, Hay, and V-8 agars (Simmons
and Roberts 1993, p. 136) in plastic Y-plates at ca. 22C
under a 10/14hr, cool-white fluorescent light/dark cycle;
microscopic details of material mounted in lactic acid are
recorded from observations made at 195X and 780X.
106. Alternaria infectoria Simmons (1986) Fig. 84
Cultures ex type of A. infectoria (EGS 27-193; Alt.T&V
23, Simmons 1986) sporulate abundantly as rather open, bushy
clumps of geniculate and branching secondary conidiophores
produced by and separated by secondary conidia. The simplest
form of branching complex originates from the pseudorostrate
tip of a single primary conidium borne at the tip of a simple
primary conidiophore. The initial (i.e., primary) conidio-
phore may remain simple or it may become elongate and vari-
ously geniculate at several successive conidiogenous loci,
yielding a fairly dense clump of simple sporulation struc-
tures. It may arise directly from the agar surface or
originate as a morphologically distinct branch of an aerial
hypha.
Individual sporulation clumps of typical A. infectoria,
when viewed at 50X, appear relatively open and fragile due
to the rather narrow dimensions of the conidia and the domi-
nating framework of secondary conidiophores, in contrast to
the visibly more robust character of these structures in the
Argentina wheat isolate (Alt.T&V 107).
Elements of Fig. 84 emphasize the appearance of the
secondary conidiophores, which can be of any length up to ca.
125 ym. The secondary conidiophore that developes directly
from the first-formed conidium usually is much longer than
ones produced later in the branching system, often equalling
at least half the height of its sporulation clump. The low-
magnification drawings illustrate sizes of conidia and
conidiophores as they develop in young clumps or in major
branches of more complex ones.
412
Fig. 84. Alternaria infectoria: sporulation habit (conidium
bodies opaque, conidiophores clear; line = 50 ym); primary
conidiophores, conidia with secondary conidiophores, and
terminal conidia (line = 50 yum); from ex-type culture on PCA.
413
All chain formation in Alternaria depends on an alter-
nation of conidia and secondary conidiophores (pseudorostra).
The pattern of prominent secondary conidiophore elaboration
and interspersed conidia is a conspicuous character of the
A. infectoria group, and this readily observed arrangement
is being used here in an attempt to segregate this group from
among numerous somewhat similar taxa for systematic purposes.
In contrast, for example, relatively inconspicuous secondary
conidiophores (tending toward 1-2 short cells) are the norm
in the similarly bushy sporulation clumps typical of the A.
alternata complex of species.
107. Alternaria triticimaculans Simmons & Perellé, sp. nov.
Fig. 185) & 66
Etym.: L., triticum, wheat + maculans, spotting
Ex culturis in agaro PCA descripta. Coloniae mycelio
epiagaro laxe lanoso vel araneoso copiose conidiogenae.
Sporulatio in catenis ramosis conidiorum 30-40++ aggregata.
Conidiophora elongata, ramosa, geniculata, ca. 130+ x 3-4ym.
Conidia ovoidea vel longiovoidea vel ellipsoidea, ad 40-45
x 9-1l “wm, erostrata, longipseudorostratescentia, sublevia
vel aspera, sordide fulva, 6-7 transverse et parce longitu-
dine septata. Habitatio typi in folio Triticum aestivum L.,
La Plata, Argentina. Holotypus: pars ex cultura EGS 41-050
(EGS isol. ex specimine A. Perellod leg., 23-II-1993 misso)
desiccata et in BPI conservata.
Spore samples from leaf surfaces contain numerous well-
developed, beakless, long-ovoid and long-ellipsoid conidia
along with roughly equal numbers of similar conidia that have
developed an apical secondary conidiophore about half as long
as the spore body. Beakless conidia have a hemispheric or
short conical apical cell. Conidia that may appear to have
a gradually narrowing beak are, in fact, in the process of
generating a pseudorostrum. The site of change from body to
beak usually can be seen as a slight break in wall contour,
color, and ornamentation. Beakless conidia and conidium
bodies reach a size range of ca. 30-60 x 11-14 yum and are a
dilute dull yellowish brown, smooth to moderately rough, and
5-10 transeptate with 1-2 longisepta in 3-6 of the transverse
segments. The approximately equal percentages of beakless and
pseudorostrate conidia might lead to inference that this is
a taxon that is limited to production of short chains of 2-3
conidia in nature. Perhaps; but I consider it more likely
that restricted sporulation on the field specimens I have
CT eee
Fig. 85. Alternaria triticimaculans: conidia from field
specimen source of holotype (line = 50 wm).
415
seen was governed by ecological factors (humidity, tempera-
ture, light) that permitted only one or two spurts of
sporulation before conditions changed. In any case it is an
observation worth recording, for the sporulation pattern
itself, though not conidium morphology, is much different in
culture than has been seen in nature.
Typical conidia from the host surface germinate readily
and, within 2-4d under my routine conditions, yield abundant-
ly sporulating colonies on PCA, Hay, and V-8 agars.
Young colonies (5-/7d) in PCA plates are loosely lanose
to arachnoid. They have very abundant sporulation from co-
nidiophores that originate at the agar surface or from aerial
hyphae. Primary conidiophores are simple with a single
apical conidiogenous locus, commonly ca. 75 x 3 ym; or they
may become longer (130 wm or more), variously branched, and
successively geniculate at a few conidiogenous loci. First-
formed conidia tend to be narrowly elliptical or ovoid and
erostrate until production of apical secondary conidiophores
begins. Primary conidia are ca. 40-45 x 9-11 wm, almost
smooth to visibly but not heavily ornamented, dilute dull
yellowish brown, with 6-7 transepta and either no longisepta
or only 1 in 1-3 of the transverse divisions. A solitary
elongating secondary conidiophore usually develops from each
primary conidium apex. However, it is not uncommon in this
taxon for a group of 2-3 subparallel pseudorostra to have a
common base cell or for a pair of pseudorostrum branches to
originate in a spreading dichotomous manner just below a
terminal conidiogenous locus.
Sporulation clumps have much the same pattern of
seemingly unlimited complexity as that seen in A. infectoria,
with series of secondary conidia separated by pseudorostra
of various lengths and numbers of geniculations (conidiogen-
ous loci). The open, bushy aspect of sporulation clumps in
A. triticimaculans, as with that of A. infectoria, depends
almost entirely on pseudorostrum branching and bending. Only
rarely does a conidium itself generate a branch by producing
a lateral or basal conidiophore. Single sporulation clumps
commonly have 30-40++ conidia. Secondary conidia are similar
to primary conidia in size, color, septation, and ornamenta-
tion; both types are smaller than fully developed conidia
found on the natural substrate.
Sporulation clumps of A. triticimaculans appear (at
50X) more robust and more strongly branched than do those of
A. infectoria. This appearance derives from the overall
416
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Fig. 86% Alternaria triticimaculans: sporulation habit
(conidium bodies opaque, conidiophores clear; line = 50 wm);
primary conidiophores, conidia with secondary conidiophores,
terminal conidia (line = 50 ym); from ex-type culture on PCA.
417
greater size of conidia and more conspicuous secondary
conidiophores of A. triticimaculans as compared with those
Oren uRreccoria.,
Natural growths of A. triticimaculans are known at
present only from specimens on infected wheat leaves that
originated with Analfa Perell6d, Universidad Nacional de La
Plata, La Plata, Bs. As., Argentina. Several 1l-conidium
isolates were obtained from one such specimen; one of these,
EGS 41-050, has been used in these comparisons and is cited
in dried form as holotype of the species.
108. Alternaria oregonensis Simmons, sp. nov. Big. 6/
Ex culturis in agaro PCA descripta. Coloniae mycelio
epiagaro vel aerie araneoso dense conidiogenae. Sporulatio
in catenis ramosis conidiorum 100+ aggregata. Conidiophora
primigena elongata, geniculata, ca. 100-150 x 3-4 ym;
conidiophora secundaria geniculata, ca. 20-60 x 3 ym.
Conidia primigena plerumque longe ellipsoidea, ca. 35-60 x
10-15 4m, 7-11 transverse septata et 1-4 longitudine septata,
dilute sordide fulva, pseudorostratescentia; conidia secunda-
ria ovoidea, obconica vel ellipsoidea, ca. 23-33 x 8-14 ym,
levia vel conspicue aspera, sordide fulva, 5-7 transeptata
et parce longiseptata. Habitatio typi in folio Triticum
aestivum L., Oregon, U.S.A. Holotypus: pars ex cultura EGS
29-194 desiccata et in BPI conservata.
Colonies 5-7 days old on PCA sporulate abundantly in
dense, often crowded, bushy clumps directly on the agar
surface or in more open clumps on loosely arachnoid aerial
mycelium, essentially all of which is conidiophorogenous.
Aerial clumps derived from a single primary conidiophore may
consist of a few to a few dozen conidia. Surface clumps
commonly sporulate so abundantly that an estimate of 100+
conidia per clump is reasonable.
Primary conidiophores 100-150 x 3-4ym readily become
geniculate at 1-5+ successive conidiogenous loci. First-
formed conidia are predominantly long-ellipsoid, although
some short-ovoid primary conidia also are present. Conidium
bodies tend to be in a size range of ca. 35-60 x 10-15 um
when apical secondary conidiophores are produced. Such
conidia have 7-11 transepta and usually 1 longiseptum in each
of 1-4 of the transverse segments; 3-4 of the earliest-formed
transepta and major longisepta become slightly thicker and
418
darker than younger septa. Secondary conidiophores become
geniculate and conidiogenous at 1-4+ loci; or occasionally
they develop in pairs from the apical cell of a conidium or
arise laterally from the basal cell. They are 20-60 x 3 yum,
longest on conidia at or near the base of a sporulation clump
and usually shorter, even only 1-2-celled, in the distal
portions of the branching chains. Beakless terminal conidia
in a well-developed clump are ovoid, obconical or ellipsoid.
Secondary conidia are ca. 23-33 x 8-14ym, smooth to conspic-
uously rough, dilute to slightly darker dull yellowish brown,
with 5-7 transepta and usually one longiseptum in 1-5 of the
transverse segments.
Surface sporulation clumps in A. oregonensis cultures
on PCA originate in long, narrow, abundantly transeptate
primary and early secondary conidia, which generate chains
of large numbers of relatively small, short-beaked conidia,
resulting in dense bushy clumps. In comparison, surface
clumps of A. triticimaculans originate in shorter, narrow
primary conidia with fewer and less heavily pigmented septa;
these conidia generate an open, branching system of pseudo-
rostrate and beakless conidia which in general are compara-
tively large and robust when fully developed. Sporulation
clumps of A. infectoria, in contrast, are open and almost
delicate in appearance due to the dominance of long narrow
secondary conidiophores and the abundance of relatively small
secondary conidia. The sporulation mass, composed of
thousands of individual clumps, is dense and, to the unaided
eye, dull black in PCA cultures of each of the three taxa.
The isolate (EGS 29-194) upon which this discussion of
A. oregonensis and its dried-culture holotype are based
originated in the laboratory of C. L. Leach, Oregon State
University, Corvallis. The source was a leafspot of wheat
noted in the spring of 1970. Positive pathogenicity was
demonstrated at that time by R. L. Powelson (personal commu-
nication), who sent additional isolates of the taxon from
leafspot of wheat in Oregon in 1972.
109. Alternaria metachromatica Simmons, sp. nov. Fig. 88
Etym.: Gr. meta-, change; chromatikos, of color
Ex culturis in agaro PCA descripta. Coloniae mycelio
epiagaro vel aerie araneoso copiose conidiogenae, griseocae-
ruleae, fuscantes. Sporulatio in catenis ramosis conidiorum
multorum aggregata. Conidiophora primigena simplicia;
Eig Clic Alternaria oregonensis: primary conidiophores,
conidia with secondary conidiophores, and terminal conidia
(line = 50 wm); from ex-type culture on PCA.
420
secundaria geniculata, 1-4 conidiogena, 25-40 yum longa.
Conidia primigena et secundaria ovoidea vel ellipsoidea,
distincte scabra, 3-5 transverse et 0-1 longitudine septata,
pseudorostrata. Conidia tertiaria et in catenis sequentibus
ovoidea vel sphaerica, 0-3 transeptata, brevipseudorostrata
vel erostrata, scabra. Conidia, conidiophora et hyphae
aeriae dilute caeruleae in aqua, rosae in acido lactico.
Habitatio typi in semine Triticum aestivum L., South Austra-
lia, Australia. Holotypus: pars ex cultura EGS 38-132
desiccata et in BPI conservata.
Colonies 3-5 days old on PCA sporulate abundantly in
dense bushy clumps directly on the agar surface and in more
open but eventually dense clumps on loosely arachnoid aerial
mycelium, essentially all of which is conidiophorogenous.
Young sporulating colonies viewed in either incident or
transmitted light have a definite blue color, near Light Sky
Blue and Sky Grey (Ridgway 1912). The color becomes darker
and duller as the colony ages to 8-10 days. Surface clumps
and aging aerial clumps are so abundantly sporogenous in a
compact bush of branching chains that an estimate of several
hundred conidia per clump can be made.
Primary conidiophores tend to be simple or meagerly
branched, not abundantly or conspicuously elongate and multi-
geniculate. First-formed conidia are ovoid or ellipsoid with
distinctly rough outer walls, 3-5 transepta and no longisepta
or at most one in 1-2 of the transverse segments, ca. 25-35
x 5-10 yum. Primary conidia and a small percentage of second-
ary conidia in developing chains produce relatively long
apical secondary conidiophores ca. 25-40 yum long. Secondary
conidiophores often develop as strongly geniculate branches
that are conidiogenous at 1-4+ loci. Primary and secondary
conidia of A. metachromatica produce relatively long pseudo-
rostra during early development of chains and, therefore, can
be considered along with other taxa of the A. infectoria
species group. However, subsequent distal sporulation in
chains is so rapid that most conidia do not develop beyond
a stage of 1 or 2 to 4 cells, rarely have a longitudinal
septum, and have only a short l-celled pseudorostrum or
simply a morphologically erostrate but conidiogenous apex.
Conidia in the distal half of chains are ca. 4-18 x 4-8 wm.
Most conidia have slightly to heavily ornamented walls.
Conidia, conidiophores, and aerial hyphae remain blue when
mounted in water. They turn pink instantly when mounted in
lactic acid. The color appears to be concentrated in the
walls and septa of conidia.
421
ses
-<-
Fig. 88. Alternaria metachromatica: primary conidiophores,
primary and secondary conidia with secondary conidiophores,
and terminal chains of conidia (line = 50 wm); from ex-type
culture on PCA.
422
The isolate (EGS 38-132) of A. metachromatica upon
which this discussion and the dried-culture holotype are
based was received in October 1985 via IMI 295682 as an
"Alternaria alternata (albino strain)," isolated from seed
of Triticum aestivum grown in South Australia. The strain
is unique in that, except for truly hyaline variants, I have
never seen another isolate of Alternaria with colors other
than shades of brown or mixtures tending toward greenish or
yellowish browns.
Hyaline, totally non-pigmented ("albino") strains of
Alternaria occasionally arise spontaneously in otherwise
ordinary dematiaceous colonies. They can be isolated easily
and usually remain hyaline through periods of routine serial
transfer and conservation. A. metachromatica, however, is
not such a hyaline entity, for its pink color in simple
lactic-acid-based mounting media and its blue color in
culture and in water, though paler than the usual Alternaria
browns, are not lost under these conditions. Nevertheless,
I suspect that A. metachromatica, bright hued in the midst
of innumerable dark-spored taxa, may prove eventually to have
a dematiaceous analogue. The taxon should be recognizable
in culture on the basis of its A. infectoria group affinity,
viz., the initial decidedly pseudorostrate chain production,
plus the enormous numbers of small, poorly septate conidia
that make up the bulk of each large sporulation clump.
110. Alternaria triticina Prasada & Prabhu (1963) Fig. 89
Original material of A. triticina exists in the form
of a field specimen holotype collected in India on leaves of
Triticum aestivum (HCIO 27619; EGS 29-131) and a derivative
isolate (Indian Type Culture Collection (Fungi) no. 1186
G.C.; EGS 17-061). The isolate remains in excellent sporu-
lation condition 30 years after its initial isolation.
Simmons (1981, Alt.T&V 5) stressed the taxonomic
importance of characteristics of conidia produced in culture
and wondered whether or not the M. B. Ellis treatment of the
taxon (1976) presented an accurate picture of the beak and
ornamentation of conidia. It is obvious to me now, after
restudying the material, that Ellis prepared his description
from authenticated field material without reference to a
culture. In any case, the kinds of beaks illustrated by
Ellis and mentioned in Prasada and Prabhu are secondary
conidiophores produced from the blunt or broadly tapered apex
sporulation habit (conidium
= 50 wm); primary
and
Alternaria triticina:
bodies opaque, conidiophores clear; line
Fig. 89.
conidiophores, conidia with secondary conidiophores,
50 wm); from ex-type culture on PCA.
terminal conidia (line
424
of initially erostrate conidia. Walls of conidia present in
the type, reported as smooth by Ellis and by the original
authors, rarely have any visible roughness. In characters
other than beak, the original description, that of Ellis, and
the results of my examinations of the type specimen are
essentially the same.
Primary conidia of A. triticina in the field specimen
are at first narrow-ovoid, becoming broad-ovoid or broad-
ellipsoid, beakless or broadly tapered, 50-80 x 20-25 yum,
sometimes producing a secondary conidiophore 20-40 x 2-3 yum.
Conidia have up to 8-9 complete or partial transverse septa
and 1-3 longitudinal septa in each of the broadest transverse
segments, are yellowish brown, and essentially smooth.
Colonies 5-7 days old on PCA sporulate abundantly as
distinct clumps of only (1-)2-6 conidia borne on primary
conidiophores that arise in large numbers directly from the
agar surface as well as from sparse arachnoid aerial hyphae.
Conidia are ovoid to broadly ovoid or ellipsoid, erostrate,
often somewhat inequilateral, up to ca. 40-60 x 14-22 yum,
dilute tawny brown, conspicuously punctate to verrucose, and
up to 6-8 transeptate with 1-2 longisepta in each of 4-5 of
the transverse segments. The pseudorostrum of primary
conidia often becomes 1l-geniculate and bears short pseudo-
rostrate secondary conidia and erostrate terminal conidia.
The taxon is very distinctive, both in conidium
morphology and particularly in the very restricted number of
conidia per sporulation unit. The primary conidium in an A.
triticina sporulation clump usually bears a well-developed
geniculate pseudorostrum which, arguably, might permit its
consideration as a rudimentary taxon of the A. infectoria
group. However, it cannot be mistaken readily as any member
of the A. infectoria group discussed above. No other
usefully stable isolate that has reached me with a suggested
determination of A. triticina (usually among miscellaneous
isolates from wheat sent for opinion) has proved identifiable
with the ex-type isolate, although there is abundant liter-
ature on the biology and pathology of the species.
111. Alternaria triticicola V. G. Rao (1964) Figes. 20
Authentic Rao material of A. triticicola is known from
portions of field material on Triticum aestivum collected in
India (HCIO 28262; EGS 29-132 and IMI 107601; EGS 24-145).
Fig. 90. Alternaria triticicola: primary conidiophores,
primary conidia with apical secondary conidiophores, and
terminal conidia from field specimen holotype (line = 50pm).
426
These possibly are parts of the same collection but are not
necessarily so (on evidence of specimen labels compared with
type description data).
Conidia of A. triticicola in the field material are at
first narrow-ellipsoid or long narrow-ovoid, ca. 60-70 x 10-
15 wm, becoming broad-ovoid or broad-ellipsoid and broadly
tapered, 80-160 x 18-25 wm, often producing an apical
secondary conidiophore ca. 15-60 x 4-5ym. Conidia have as
many as 15-20 complete or partial transepta and 1-4 longi-
septa in most of the transverse segments; they are distinctly
constricted at major transepta, clear tawny brown, and
smooth.
The morphology of young conidia and of large conidia
before pseudorostrum production, and the comparatively large
size of fully developed conidia aid in differentiating A.
triticicola from A. triticina (Alt.T&V 110). The proportion
of pseudorostrate conidia to erostrate (i.e., terminal)
conidia suggests that chain formation of several units (2-3
according to the type description) is characteristic of the
taxon in nature.
ACKNOWLEDGEMENTS
My thanks continue to reviewer Dr. Mary E. Palm, who
never lapses in catching mine. Her reviews and editorial
comments on my manuscripts are always specific, valuable, and
much appreciated.
The Alternaria taxon received from Ms. Analfa Perellé
in the course of her study of a related wheat disease in
Argentina can be expected to be included in her future
phytopathology papers. The sources of other Alternaria taxa
from wheat are acknowledged gratefully: Australia (leg.
incogn.) via IMI, International Mycological Institute, Egham,
U.K.; India, HCIO, Indian Agricultural Research Institute,
New Delhi; Oregon, U.S.A., C. L. Leach and R. L. Powelson.
LITERATURE CITED
BLL is) Mae Lo TGe More Dematiaceous Hyphomycetes.
Commonwealth Mycological Institute, Kew. 507 pp.
Holmgren, P. K., N. H. Holmgren, and,L. C. Barnett. 1990.
Index Herbariorum Part I: The Herbaria of the World.
427
8th ed. New York Botanical Garden, Bronx. 693 pp.
Prasada, R., and A. S. Prabhu. 1963. Leaf blight of wheat
caused by a new species of Alternaria. Indian Phyto-
Path fo 962) 72924293"
Rao, V. G. 1964. An undescribed species of Alternaria on
wheat from India. Mycopath. Mycol. Appl. 23: 311-313.
Ridgway, R. 1912. Color Standards and Color Nomenclature.
Ridgway (as publisher), Washington, D.C. 44 pp.+ 53
plates.
Simmons, E. G. 1981. Alternaria themes and variations [1-
6]. Mycotaxon 13: 16-34.
Simmons, E. G. 1986. Alternaria themes and variations (22-
26). .Mycotaxon 25;..28/- 308.
Simmons, E. G., and R. G. Roberts. 1993. Alternaria themes
and variations (73). Mycotaxon 48: 109-140.
MY COTAXON
Volume L, pp. 429-433 January-March 1994
MENISPORA CONVOLUTA, A NEW DEMATIACEOUS HYPHOMYCETE FROM
CENTRAL ITALY
DARIO LUNGHINI
Laboratorio di Micologia, Dipartimento di Biologia
Vegetale, Universita degli Studi di Roma "La Sapienza",
Largo Cristina di Svezia 24, 00165 Roma, Italy.
SUMMARY
A new dematiaceous Hyphomycete species, Menispora
convoluta, found on juniper wood in central Italy, is
described and illustrated.
Keywords: Hyphomycetes, Italy, Menispora convoluta.
During a mycological investigation carried out in the
Castelporziano Presidential Estate, near Rome (Lunghini &
Quadraccia, 1990; Quadraccia & Lunghini, 1990), the
author found a species of Menispora Pers. on juniper
wood, showing peculiar characteristics, but the scarcity
and bad condition of the specimen did not allow a
comprehensive study. A subsequent finding of the same
species, also on juniper wood, near Campo Cecina (Alpi
Apuane, Massa Carrara), made it possible to conclude the
work and describe the following new species.
Menispora convoluta Lunghini sp. nov.
Coloniae effusae, haud constantes, lanatae villosaeve,
umbrinae, conidiophora et setas steriles ferentes.
Mycelium partim superficiale, partim immersum, ex hyphis
septatis, ramosis, a subhyalino ad brunneum colorem
variantibus, 1.5-4 um crassis compositum.
Setae steriles amplitudine et colore conidiophoris
similes sed raro- ramis praeditae. Conidiophora
setiformia, anguste conferta, luteo-brunnea, sursum
430
leviter attenuata, usque ad 950 ym longa et ad basim
3.5-5 jm crassa, septata, vulgo enormiter ramosa.
Conidiophora exeunt in cellulas conidiogenas vel in
longas setas sursum connvolutas et anguste inter se
complicatas.
Cellulae conidiogenae phialidicae, subcylindricae,
clavulatae, in parte inferiore rectae vel raro
recurvatae, pallidiores quam conidiophori stipes, 24-48
pm longae et 3.5-5 wm crassae, repente obtruncatae, ad
apicem conspicue recurvatae, sessiles aut praeditae brevi
pediculo 2-4 cellularum instructo, collulo inconspicuo
vel nullo praeditae. Phialides terminales rursus gignere
possunt seu alteram phialidem seu setam convolutam.
Phialoconidia in capitulo mucido producta, hyalina,
continua, subcylindrica vel cylindrico-ovata, leviter
curvata, ad basim acutioria quam ad apicem, 17-23 x
3.5-4.2 pm, utrinque setula 5.5-9.5 wm longa praedita.
In ligno putrido Juniperi communis L., Campo Cecina
(Alpi Apuane, Massa Carrara), Italy, Lunghini 5-11-1990,
HOLOTYPUS: ROHB 140.
Additional specimen: on rotten wood of Juniperus
oxycedrus L. ssp. macrocarpa (Sibth. et Sm.) Ball, Tenuta
Presidenziale di Castelporziano, Roma, Italy, Lunghini
AZ 21989 2
Colonies effuse, irregular, woolly to hairy, DARK
BROWN, bearing noumerous conidiophores and sterile setae.
Mycelium partly superficial, partly immersed, composed of
septate, branched, subhyaline to brown, 1.5-4 pm wide
hyphae.
Sterile setae similar in colour, size and appearance
to conidiophores, but rarely branched. Conidiophores
setiform, tightly crowded, YELLOWISH BROWN, paler towards
the apex, up to 950 wpm long, 3.5-5 pm wide near the
swollen base, gradually tapering to 2 wpm near the
rounded apex, septate, repeatedly and irregularly
branched. Conidiophores and their branches ending in
conidiogenous cells or distally twisted setae, which
become entangled with each other, but do not usually
anastomose.
Conidiogenous cells subcylindrical, clavulate,
straight or slightly curved in the lower part, paler than
the conidiophore stipe, 24-48 x 3.5-5 ym, abruptly
tapered and recurved at the apex, sessile or borne on
short branches of 2-4 cells. Collarette inconspicuous or
H* 4)
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Menispora convoluta Lunghini
432
absent. Terminal phialides sometimes proliferating
percurrently into new phialides or developing coiled,
seta-like extentions, that in turn can branch out,
bearing new phialides and setae. Traces of proliferation
are easily recognizable as constrictions along the
full-grown stipes and branches of the conidiophore.
Similar configurations can be found in some other
phialidic hyphomycetes; even if they are usually remnants
of percurrent proliferation, the light microscopic
observation of the constrictions suggests continuity in
the outer wall. This suggest an holoblastic mode of
regeneration and development.
Conidia aggregated in slimy balls, hyaline, aseptate,
subcylindrical or cylindrical-obovoid, slightly curved,
asymmetrical, rounded at the distal end and tapered at
the proximal end, 17-23 x 3.5-4.2 pm, bearing on either
ends a single, 5.5-9.5 pm setula. An attempt to obtain
this species in pure culture was unsucessful. No
ascocarps of a teleomorph state were found.
The described species can easily be included in the
genus Menispora Pers., mainly on the basis of the
morphology of conidiogenous cells and conidia. Among the
species ascribed to this genus, mM. convoluta presents
affinity with M. gamsii Hol.-Jech. and, to a lesser
extent, M. caesia Preuss. M. convoluta differs from these
species in having very coiled setae, more intricate
branching, dissimilar colours and dimensions and in other
minor features, but especially in the ability of the
phialides to proliferate or produce holoblastically
secondary sterile setae. Phialide proliferation is known
in other Menispora species, viz. M. uncinata S. Hughes &
B. Kendrickvand'M.!’ britannica) (M.Bu Ellis)uPoM. Kirk,© but
in these species proliferation is sympodial.
Moreover, it should be noted that in spite of the
similarity of the substrata, the two localities differ
considerably in climatic and biogeographical conditions.
Castelporziano Presidential Estate is a typical
mediterranean environment at sea level, whereas Campo
Cecina, on the Apuan Alps at about 1200 m above sea
level, is a submountain station, even if influenced by
mediterranean conditions. However, both localities can be
considered as adverse to the survival of Hyphomycetes and
the production of strongly coiled setae can be regarded
as mode of protection for the conidia.
433
The author wishes to thank Prof. W. Gams for kindly
reviewing the manuscript, Prof. M. Bigazzi (Rome) for
assistance with the Latin diagnosis and Dr. A. Pavese
(Rome) for classifying substrata. This research was
supported by a grant from M.U.R.S.T.
REFERENCES
HOLUBOVA'-JECHOVA', V. (1973). Lignicolous Hyphomycetes
from the Netherlands. Proc. Kon. Ned. Akad. Wetensch.,
Ser. 1G, 32) 297-502.
HUGHEoy) S.J. & KENDRICK, W.B. (1963). Microfungi. IX.
Menispora Persoon. Can. J. Bot., 41: 693-718.
HUGHES, S.J. & KENDRICK, W.B. (1968). New Zealand Fungi.
12. Menispora, Codinaea, Menisporopsis. N. Z. Jl Bot.
Oe S20-5./.0%
KIRK, P.M. (1985). New or interesting Microfungi. XIV.
Dematiaceous Hyphomycetes from Mt Kenia. Mycotaxon 23:
BUO-352.
LUNGHINI, D. & QUADRACCIA, L. (1990). Contributo alla
conoscenza degli Ifomiceti demaziacei della Tenuta
Presidenziale di Castelporziano (Micoflora del Lazio
III). Quaderno Accad. Naz. Lincei 264: 121-132.
QUADRACCIA, L. & LUNGHINI, D. (1990). Contributo alla
conoscenza dei macromiceti della Tenuta Presidenziale
di Castelporziano (Micoflora del Lazio II). Quaderno
Accad. Naz. Lincei 264:749-120.
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MY COTAXON
Volume L, pp. 435-440 January-March 1994
REDISPOSALS AND REDESCRIPTIONS IN
THE MONOCHAETIA - SEIRIDIUM,
PESTALOTIA - PESTALOTIOPSIS COMPLEXES.
X. PESTALOTIA GRANATT AND
PESTALOZZINA PUNICAE
T. R. NAG RAJ
Department of Biology, University of Waterloo, Waterloo,
Ontario, Canada. N2L 3G1.
and
V. MEL’NIK
Botanical Institute of the Russian Academy of Sciences,
Prof. Popov Street 2, 197 376 St. Petersburg, Russia.
Pestalotia granati is redisposed in Pestalotiopsis.
Pestalozzina punicae is considered a nomen dubium.
Husseinov (1968) gave an illustrated account of Pestalotia
granati Husseinov from living leaves of Punica granatum
collected in Azerbaijan (Commonwealth of Independent States
- former U.S.S.R.). Diagnostic features for the species were
presented as follows: 1) numerous black, epiphyllous, scattered
to aggregated acervuli; 2) clavate, rarely subclavate, 4-septate
436
conidia 17.5-20 X 5.5 um, with brown or pale brown median
cells with combined length of 11-14 tum; 3) colourless or almost
colourless apical and basal cells; 4) filiform, divergent, apical
appendages, 10-13 jim long, and 5) a basal appendage, 3-4 um
long.
Our study of the type specimen in LE 73885 leads us to
conclude that the fungus is more appropriately disposed in
Pestalotiopsis as Pestalotiopsis granati. The nomenclator and a
revised description are presented below:
Pestalotiopsis granati (Husseinov) Nag Raj & Mel’nik comb.
nov. Fig. 1.
= Pestalotia granati Husseinov, Mikol. 1. Fitopatol. 2: 512,
1968.
Foliicolous. Conidiomata stromatic, pycnidioid, predominantly
epiphyllous, scattered to gregarious, and often confluent, innate
erumpent, appearing as angular, orbicular or irregular, pale
brown or straw-coloured elevated pustules with glistening black
centre, but after dehiscence wide open with a black mass of
conidia extruded in long cirrhi, 110-360 tum wide, 70-100 um
deep, unilocular, glabrous; basal stroma and parietal tissue 10-15
uum thick, of almost colourless textura globulosa or textura
angularis. Conidiophores lining the cavity of the conidioma,
sparsely septate and branched only at the base, mostly reduced
to conidiogenous cells, colourless, smooth, invested in mucus.
Conidiogenous cells phialidic, ampulliform to lageniform,
colourless except for the almost colourless apical part, smooth,
6-15 X 2-4 [x = 10.3 X 2.6] um, with up to 4 percurrent
proliferations. Conidia fusiform, 4-septate, 20-26 X 5-6 [XK = 22.5
437
X 5.5] ym, bearing appendages; basal cell obconic with a
truncate base, almost colourless, thin- and smooth-walled, 3.5-5
[x = 4.2] um long; median cells 3, doliiform to subcylindrical,
Figure. 1. Pestalotiopsis granati ex holotype in LE 73885.
A. v.s. of a conidioma; B. Conidiogenous cells with
developing conidia; C. Mature conidia.
438
with rugose walls, brown but often with the two upper cells
darker than the lower cell, together 13-17 [k = 14.5] um long
(second cell from the base 4.5-6.5 [x = 5.3] tm; third cell 4.5
(-5) [K = 4.5] tum; fourth cell 4-5 [k = 4.6] um); apical cell
conical, almost colourless, thin- and smooth-walled, 3-4.5 [X =
3.8] [um long; appendages tubular, unbranched, attenuated,
flexuous; appendages on the apical cell 2-3, one of them axial
and the others abaxial inserted at different loci, more or less
proximal to the base of the axial appendage, 7-13(-16) [k = 10]
tum long; appendage on the basal cell, when present, single,
unbranched, centric, up to 4 [x = 2.3] um long; mean conidium
length/width ratio = 4.1:1.
Habitat: On living leaves of Punica granatum.
Specimen examined: LE 73885 [Type], pagum Kanar-Mesha,
Dist. Lenkoran, Transcaucasia, Azerbaijan, 15.VII.1967,
E.S.Husseinov.
Known distribution: Azerbaijan (Commonwealth of
Independent States - former U.S.S.R.)
Teleomorph: Unknown.
Pestalotia jodhpurensis Bilgrami & Purohit, the only other
species known to occur on P. granatum in India, belongs in
Pestalotiopsis but is different from Pestalotiopsis granati in its
conidium morphology and dimensions. P. granati is associated
with greyish, round or irregular leaf spots, 1-1.3 cm wide.
Husseinov (1970) published the binomial Pestalozzina
punicae for a fungus occurring on living leaves of Punica
granatum [Evlach, Azerbaijan (Commonwealth of Independent
States - former U.S.S.R.), 10.[X.1968, E.Husseinov]. The fungus,
observed on small, brown, round or elongate spots, was reported
439
to have: 1) amphigenous, small, brown acervuli; 2) cylindrical
or oblong-clavate, 4-septate, colourless (yellowish in mass)
conidia 15.6-23.3 X 4-5 um, and 3) 2-3, filiform appendages at
the apex, 5-13 X 1-1.5 tm and 4) a basal appendage 2.9-3.7 um
long. LE was the only reported repository for the type specimen;
however, the specimen does not exist in that herbarium. The
reported conidial dimensions fall well within the limits for
Pestalotiopsis granati, but the published description and
illustration of the fungus cast some doubts on its affinity with
Pestalotiopsis. It is more likely to be a species of Zetiasplozna
Nag Raj (Nag Raj 1993), but its true affinities cannot be
ascertained at this time. We prefer to relegate the binomial to the
status of a nomen dubium.
Acknowledgments
We are grateful to Prof. Bryce Kendrick, Dept. of Biology,
University of Waterloo, and Dr. Amy Rossman, Research
Leader, Systematic Botany and Mycology Laboratory,
U.S.Department of Agriculture, Beltsville, Maryland, U.S.A., for
Critical reviews of the manuscript. One of us (T.R.N.)
acknowledges financial support from the Natural Sciences and
Engineering Research Council of Canada in the form of an
operating research grant to Prof. Kendrick.
References
Husseinov, E.S. 1968. [New species of fungi from Azerbaijan].
Mikol. i. Fitopatol. 2(6): 512-514.
Husseinov, E.S. 1970. [New species of fungi on fruit trees in
Azerbaijan]. Mikol. i. Fitopatol. 4(1): 73-75.
440
Nag Raj, T.R. 1993. Coelomycetous anamorphs with appendage-
bearing conidia. 1101 pp. Mycologue Publications, Waterloo,
Ontario, Canada.
MY COTAXON
Volume L, pp. 441-458 January-March 1994
CYLINDROCLADIUM NAVICULATUM SP. NOV., AND TWO NEW
VESICULATE HYPHOMYCETE GENERA, FALCOCLADIUM AND
VESICLADIELLA
P.W. CROUS
Department of Plant Pathology, University of Stellenbosch, 7600
Stellenbosch, South Africa
M.J. WINGFIELD
Department of Microbiology and Biochemistry, University of the Orange
Free State, P.O. Box 339, 9300 Bloemfontein, South Africa
A.C. ALFENAS AND S.F. SILVEIRA
Departemento de Fitopatologia, Universidade Federal de Vigosa, 36570
Vicgosa, Brazil
ABSTRACT
Cylindrocladium naviculatum is newly described and distinguished from
other Cylindrocladium species based on its narrowly 1-septate conidia,
naviculate vesicles, and cultural characteristics. Falcocladium and
Vesicladiella are described as two new hyphomycete genera with white
sporodochial conidiomata from Eucalyptus and Araucaria leaf litter
respectively. Both genera are characterized by having thick-walled, non-
septate stipe extensions that terminate in thin-walled vesicles.
Falcocladium has appendaged, falcate conidia, whereas Vesicladiella has
aseptate, cylindrical conidia. These genera are also distinguished from
others such as Cylindrodendrum, Cylindrocladium and Cylindrocladiella by
the morphology of their conidiomata and stipe extensions.
During collections of hyphomycete specimens from Brazil, a species of
Cylindrocladium Morgan with naviculate vesicles was isolated from soil in
Manaus, Amazonas, while a vesiculate hyphomycete with sporodochial
conidiomata was isolated from Eucalyptus leaf litter at Aracruz in the
State of Espirito Santo. The latter collection resembles the genera
Cylindrocladium and Cylindrocladiella Boesewinkel in having hyaline stipe
442
extensions terminating in well differentiated vesicles. Terminal vesicles
and stipe extensions are, however, also found in genera such as
Cylindrodendrum Bonorden and Pulvinotrichum Gamundi, Arambarri &
Giaiotti. The aim of this study was to compare the Cylindrocladium
isolate with all other taxa presently known in this genus. A comparison
was also made of the morphology of the hyphomycete from Eucalyptus
with that of Cylindrodendrum and Pulvinotrichum in order to suitably
delineate these taxa based on their general morphology and cultural
characteristics.
MATERIALS AND METHODS
Morphology
Cultures derived from single conidia were plated onto carnation-leaf agar
(CLA) (Crous, Phillips & Wingfield, 1992), incubated at 25°C under
near-ultraviolet light, and examined after 7 days in the case of the
Cylindrocladium sp., and 14 days for the hyphomycete from Eucalyptus.
Only material occurring on carnation leaves was examined. Mounts were
prepared in lactophenol cotton blue. All measurements were made
under the (100 x) oil-immersion objective.
The Cylindrocladium species (CPC 627-629) had 1-septate conidia
and narrow, tapering vesicles, suggesting that it should be compared with
other 1-septate species with similar vesicle morphologies such as C.
avesiculatum Gill, Alfieri & Sobers (ATCC 38226), C. candelabrum
Viegas (PPRI 4153), C. clavatum Hodges & May (PPRI 3994), C. gracile
(Bugn.) Boesewinkel (PC 551197), C. hawksworthii Peerally (MUCL
30866), C. pteridis Wolf (PPRI 4157) and C. scoparium Morgan (ATCC
46300).
The fungus isolated from Eucalyptus leaf litter (CPC 589)
produced hyaline sporodochia with several stipe extensions and terminal
vesicles, suggesting a similarity to genera such as Cylindrocladium,
Cylindrocladiella, Cylindrodendrum, Uncigera and Pulvinotrichum.
Cultural characteristics
Growth studies: To determine the maximum radial growth of species in
culture, agar plugs (3 mm diam) from the periphery of young actively
growing colonies of each fungus were plated at the centre of malt-extract
agar (MEA) (20 g Oxoid malt extract, 15 g Difco agar, 1000 ml H7O)
plates, and incubated at 25°C for 1 day to ensure active growth. Growth
after 1 day was noted, and thereafter, plates were placed in incubators at
443
the temperatures under consideration. Isolates were incubated in the
dark at 5, 8, 10, 15, 20, 25, 30, 33 and 35°C with three replicate plates of
each isolate at each temperature. Growth was assessed after 6 days for
the Cylindrocladium spp., and 14 days for the hyphomycete from
Eucalyptus. Average growth was calculated from four radial
measurements from each of the three plates.
Chlamydospores and colony colour: Colony colour and chlamydospore
formation was determined at 25°C in the dark after 6 days for the
Cylindrocladium spp., and after 14 days for the hyphomycete from
Eucalyptus. Colour designations used were those of Rayner (1970).
RESULTS AND DISCUSSION
Cylindrocladium sp. from soil:
The Cylindrocladium species (CPC 627-629) collected in Manaus had
naviculate vesicles, which were similar to those of young, developing
vesicles of C. candelabrum (Crous, 1992). These two species could,
however, be distinguished by the characteristic apical taper of the
vesicles, the narrower conidia, moderate chlamydospore and slight
microsclerotium formation in the fungus under investigation (Table 1).
Other criteria that appear to be unique to this species are the frequent
multiple-stipes occurring in a conidiophore, as well as the very wide stipe
diameter observed just below the vesicle (Table 1). This species can
easily be distinguished from all other 1-septate species of Cylindrocladium
(Crous, 1992), and we therefore describe it here as new.
Cylindrocladium naviculatum Crous & Wingfield sp. nov. Figs 1, 5-
8.
Etym.: named after its characteristic naviculate vesicles.
Macroconidiophora. Fibra septata, hyalina, terminata vesica naviculare,
(6-)7.5(-11) wm diam; stipites (150-)180(-200) wm longi. Rami
conidiophori: rami primarii non-septati vel raro 1-septati,
(12-)18.5(-25) x (4-)4.5(-6) wm; rami secundarii non-septati,
(10-)12(-15) x (4-)4.5(-6) wm; rami tertiani non-septati,
(10-)12(-15) x (4-)4.5(-6) wm; rami quartani non-septati,
(10-)11(-14) x 4(-5) um. Phialides exorientes ex extremitatibus
ramorum, 2-4 congregatae; phialides doliiformes vel reniformes,
hyalinae, (9-)11.5(-15) x (3-)3.5(-4) wm. Conidia cylindrica, hyalina,
1-septata utrisque extremitatibus rotundatis, (40-)43(-55) x
(3-)3.5(-4) wm. Microconidiophora ignota. Teleomorpha ignota.
add
Colonies (bottom) tawney 13’i (Rayner, 1970) on MEA, obtaining a
radius of 28 mm after 6 days at 25°C in the dark. Chlamydospores in
moderate numbers, forming fine chains with sparse microsclerotia.
Temperature requirements for growth: minimum temperature above 5S°C,
maximum temperature above 35°C, optimum temperature 25°C.
Macroconidiophores. Stipe extension septate, hyaline, terminating in a
naviculate vesicle, (6-)7.5(-11) wm diam; stipes (150-)180(-200) um
long. Conidiophore branches: primary branches non-septate or rarely
1-septate, (12-)18.5(-25) x (4-)4.5(-6) um; secondary branches
non-septate, (10-)12(-15) x (4-)4.5(-6) wm; tertiary branches
non-septate, (10-)12(-15) x (4-)4.5(-6) wm; quaternary branches
non-septate, (10-)11(-14) x 4(-5) wm. Phialides arise from the ends of
branches, in groups of 2, 3 or 4; phialides doliiform to reniform, hyaline,
(9-)11.5(-15) x (3-)3.5(-4) wm. Conidia cylindrical, hyaline,
1-septate rounded at both ends, (40-)43(-55) x (3-)3.5(-4) wm.
Microconidiophores not present. Teleomorph not present.
Holotype: Brazil: Amazonas State, Manaus, soil, M.J. Wingfield, Apr.
1993, PREM 51542.
Cultures examined: Brazil: Amazonas State, Manaus, soil, M.J.
Wingfield, Apr. 1993, CPC 627 (culture of type); other collections from
the same location, CPC 628, 629.
Hyphomycete from Eucalyptus leaf litter:
The hyaline hyphomycete collected from Eucalyptus leaf litter (CPC 589)
grew and sporulated profusely in culture. Numerous thick-walled, dark
brown chlamydospores were common, and were also characteristic of
Cylindrocladium, Cylindrocladiella, Cylindrodendrum and Pulvinotrichum.
Conidiomata are hyaline, but frequently have light brown bases.
These structures originated either from hyphae, or from stromata
comprised of chlamydospores. Conidiomata are penicillate or
synnematal, but more frequently aggregated in sporodochia. Single
conidiophores include several branches, terminating in a whorl of short
to elongate ampulliform phialides. Each conidiophore has one to several
thick-walled, non-septate stipe extensions which terminate in thin-walled,
ellipsoidal vesicles. These stipe extensions originated randomly
throughout the conidiomata, and are not restricted to the terminal ends
of branches or phialide positions as is found in species of Cylindrocladium
Fig. 1. Cylindrocladium naviculatum. A, conidia; B, naviculate vesicles;
C, conidiophore (bar = 10 um).
446
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447
or Cylindrocladiella. The most characteristic feature of this fungus is its
falcate conidia. Conidia also have short basal appendages which occur at
the inner, shorter curve of the conidium, and inconspicious, short apical
appendages, continuous with the rest of the conidium.
Generic considerations:
Cylindrocladium and Cylindrocladiella: Arguments for separating these
genera have been presented elsewhere (Crous, 1992; Crous & Wingfield,
1993). The vesiculate fungus under discussion here is distinct from all
Cylindrocladium species (Crous, 1992) in having non-septate, thick-
walled stipe extensions, and falcate conidia (Figs 2, 3). Although the
nature of the stipe is similar to that of Cylindrocladiella spp. (Fig. 4),
these never form sporodochial conidiomata, or have falcate conidia
(Crous & Wingfield, 1993).
Cylindrodendrum, Pulvinotrichum and Uncigera: Summerbell et al. (1989)
recently compared these three genera, and concluded that they are
congeneric. Cylindrodendrum is characterized by conidiomata composed
of erect, hyaline hyphae bearing whorls of phialides. The only character
distinguishing Cylindrodendrum from Uncigera (Di Cosmo et al., 1983) is
the strongly bent apical portions of the phialides in Uncigera.
Summerbell et al. (1989), however, also observed this characteristic in
Cylindrodendrum, and thus reduced Uncigera to synonymy with it.
Furthermore, in an examination of the type specimen of Pulvinotrichum,
Summerbell et al. (1989) found the same characteristic conidiomatal and
phialide morphology to be present, thus supporting the synonymy of
Pulvinotrichum and Cylindrodendrum with the latter genus having
priority.
The nature of the stipe extension, the branching pattern of the
conidiophore, and terminal vesicle in Cylindrodendrum as illustrated by
Arambarri et al. (1981) and Summerbell et al. (1989) are distinct from
those observed in Cylindrocladium, Cylindrocladiella, as well as from
those of the hyphomycete from Eucalyptus under investigation here.
Furthermore, the distinct falcate conidia of this fungus preclude it from
being suitably accommodated in any of these genera. The new genus
Falcocladium is therefore proposed for this fungus.
Falcocladium Silveira, Alfenas, Crous & Wingfield gen. nov.
Etym.: named after its falcate conidia.
448
Conidiomata hyalina, sporodochialia vel synnematalia, vel penicillata
quum formata sint in mycelio aerio, reperta singulatim, sed saepius
aggregata, exorientia ex mycelio ipso vel ex stromate crassitunicatarum et
rufobrunnearum chlamydosporarum quum hic fungus cultus est in CLA;
stipitis extensiones hyalinae, non-septatae, crassitunicatae, exorientes ex
qualibet positione in ramo conidiomatali, vel in positione phialidis, et
plures quam una saepe repertae sunt in penicillato conidiomate eodem.
Rami conidiophori hyalini, non- vel multiseptati, usque tres series
ramorum per conidioma. Phialides hyalinae, exorientes ex extremitatibus
ramorum, congregatae 2-6, ampulliformes, collariculis inconspicuis.
Conidia hyalina, 0(-1)-septata, falcata acutis brevibus apicalibus et
basilaribus appendicibus.
Colonies dark brown on MEA with brown, thick-walled chlamydospores.
Conidiomata hyaline, sporodochial or synnematal, or penicillate when
formed on aerial mycelium, occurring singly, but more frequently
aggregated, arising directly from the mycelium, or from a stroma of thick-
walled, red-brown chlamydospores when cultured on CLA; stipe
extensions hyaline, non-septate, thick-walled, originating from any
position on a conidiomatal branch, or in the position of a phialide,
frequently with more than one occurring in the same penicillate
conidioma. Conidiophore branches, hyaline, non- to multi-septate, up to
three series of branches per conidioma. Phialides hyaline, arising from
ends of branches, in groups of 2-6, ampulliform, with inconspicious
collarettes. Conidia hyaline, 0(-1)-septate, falcate with acute, short
apical and basal appendages.
Sp. typ. F. multivesiculatum
Falcocladium multivesiculatum Silveira, Alfenas, Crous & Wingfield sp.
nov. Figs 2, 3,
9-11, 14-19.
Etym.: named after the numerous vesicles occurring on conidiomata.
Conidiomata sporodochialia vel synnematalia, vel penicillata quum
formata Sint in mycelio aerio, reperta singulatim, sed saepius aggregata,
exorientia ex mycelio ipso vel ex stromate crassitunicatarum
rufobrunnearum chlamydosporarum quum hic fungus cultus sit in CLA;
stipitis extensiones hyalinae, non-septatae, crassitunicatae,
(45-)60(-80) zm (mensurae a septo basilari usque ad acumen vesicae),
Fig. 2. Synnematal conidioma of Falcocladium multivesiculatum arising
from a stroma of chlamydospores (bar = 10 um).
449
450
exorientes ex positione qualibet in ramo conidiomatali, vel in positione
phialidis, plures quam una saepe repertae sunt in conidiomate eodem
penicillato. Rami conidiophori, rami primarii hyalini vel pallide brunnei,
0-4-septati, 11-95 x 3.5-5 wm; rami secundarii hyalini, 0-2-septati,
10-80 x 3.5-5 xm; rami tertiani hyalini, 0-2-septati, 10-50 x 3.5-5
pm. Phialides hyalinae, exorientes ex extremitatibus ramorum, 2-6
aggregatae, ampulliformes, 13-60 x 4-7 um; collaricula inconspicua.
Conidia hyalina, 0(-1)-septata, falcata, acutis brevibus apicalibus
basilaribusque appendicibus, (12-)14(-20) x 1.5(-2) um (appendicibus
exClusis); appendices basilares sitae in interiore breviore convexa
conidica flexione, 1-3 um longae, terminatae apice obtuse rotundato;
appendix apicalis continua conidio, 1.5-2 wm longa, terminata apice
obtuse rotundato.
Colonies (bottom) 13’k verona brown (Rayner, 1970) on MEA, attaining
a radius of 7 mm after 14 days at 25°C in the dark; aerial mycelium sparse,
margins irregular. Chlamydospores dark brown, arranged in chains,
10-20 wm long, 5-11 wm wide. Temperature requirements for growth,
minimum temperature above 10°, optimum temperature 20-25°,
maximum temperature below 35°. Conidiomata sporodochial or
synnematal, or penicillate when formed on aerial mycelium, occurring
singly, but more frequently aggregated, arising directly from the
mycelium, or from a stroma of thick-walled, red-brown chlamydospores
when cultured on CLA; stipe extensions hyaline, non-septate, thick-
walled, (45-)60(-80) ~m (measured from basal septum to vesicle tip),
originating from any position on a conidiomatal branch, or in the
position of a phialide, frequently with more than one occurring in the
same penicillate conidioma. Conidiophore branches, primary branches
hyaline to light brown, 0-4-septate, 11-95 x 3.5-5 um; secondary
branches hyaline, 0-2-septate, 10-80 x 3.5-5 xm; tertiary branches
hyaline, 0-2-septate, 10-50 x 3.5-5 wm. Phialides hyaline, arising from
ends of branches, in groups of 2-6, ampulliform, 13-60 x 4-7 wm;
collarettes inconspicious. Conidia hyaline, 0(-1)-septate, falcate with
acute, short apical and basal appendages, (12-)14(-20) x 1.5(-2) wm
(excluding appendages); basal appendages situated on the inner, shorter,
convex, conidial curve, 1-3 um long, terminating in bluntly rounded
apices; apical appendages continuous with conidia, 1.5-2 ~m long,
terminating in bluntly rounded apices.
Fig. 3. Penicillate conidiomata of Falcocladium multivesiculatum arising
from aerial mycelium (bar = 10 um).
Se b
452
Holotype: Brazil, Espirito Santo, Aracruz, Eucalyptus grandis leaf litter,
Silvaldo F. Silveira, Jan. 1993, PREM 51541.
Additional specimens and cultures examined: Cylindrodendrum album
Bonorden, Australia, NSW, Bellangry forest, Wilson River Reserve, on a
dead grass blade in middle of Wilson River, leg. R.V. Bandoni, det. K.
Seifert, 18 Aug. 1981, CBS 3484; Pseudomicrodochium candidum (Bres.)
de Hoog, Belgium, near Eupen, decaying leaf of Ilex aquifolium, Leg. &
det. W. Gams, 17 Oct. 1981, Netherlands, Callantsoog, dead leaf of
Quercus, leg. W. Gams, Oct. 1979, CBS (no number assigned);
Pulvinotrichum album Gamundi, Arambarri & Giaiotti, Argentina, Pla.
Quetrihue, leaves of Nothofagus dombeyi, leg. P. Benavente, 19 Oct. 1977,
LPS 40140; Pulvinotrichum capitatum Sutton, Australia, Queensland,
Boombana, M.P., leaf litter of Araucaria bidwillii, leg. B.C. Sutton & J.L.
Alcorn, 27 Aug. 1981, IMI 263277a; Cylindrocladiella elegans Crous &
Wingfield, RSA, Natal, Seven Oaks, leaf litter, leg. I. Rong, 11 Oct. 1989,
PPRI 4050 (culture); Cylindrocladium colhounii Peerally var. colhounii,
Brazil, unknown host, leg. A.C. Alfenas, 1991, PPRI 4183 (culture).
When synonymizing Uncigera and Pulvinotrichum with
Cylindrodendron Summerbell et al. (1989), noted that further collections,
cultural, as well as type specimen studies would be required to consider
possible synonymies among the species previously described in these
three genera. Unfortunately Sutton (1989) described a new species,
Pulvinotrichum capitatum Sutton in the same year. The latter collection
was placed in Pulvinotrichum due to its sporodochial conidiomata, as well
as the presence of stipe extensions with terminal vesicles as illustrated by
Arambarri et al. (1981). An examination of P. capitatum (IMI 263277a)
however, showed that in spite of several similar characteristics, this
Species cannot be accommodated in Cylindrodendrum. The conidia of P.
capitatum are cylindrical, aseptate, and carried in cylindrical packets. The
same was also found to be true for several collections of Cylindrodendrum
that we examined. Furthermore, phialides of P. capitatum could also be
Fig. 4. Falcocladium and morphologically similar genera. A, Penicillate
conidiophore of Cylindrocladiella camelliae (CPC 395); B, phialides and
conidia of Cylindrodendrum album (CBS 3484); C, phialides, conidia and
vesicles of Cylindrodendrum album var. paralion (redrawn from
Summerbell et al., 1989); D, curved phialides and conidia of
Cylindrodendrum album (type of Pulvinotrichum album, LPS 40140); E,
conidiophores with phialides, conidia and vesicles of Vesicladiella
capitatum (IMI 263277a) (bar = 10 wm).
454
accommodated within the range of variation described for
Cylindrodendrum, with some also having the characteristically abrupt
apical bend to the upper part of the phialide (Fig. 4).
Pulvinotrichum capitatum and Cylindrodendrum differ in the nature
of the stipe extension and the phialide bearing hyphae (Fig. 4). In P.
capitatum, stipe extensions are thick-walled, non-septate, with several
occurring in every conidioma, and branched at 90° to the basal hyphae.
This was never the case in Cylindrodendrum, where stipe extensions were
rare, septate, thin-walled, and rather extensions of fertile developing
hyphae with indeterminate growth. Colonies are also not as floccose as
those of Cylindrodendrum. Furthermore, in contrast to Cylindrodendrum
where the phialide bearing hyphae are erect, thin-walled, and carry
whorls of phialides, hyphae of P. capitatum are thick-walled, occur on the
substrate surface, with the phialides not arranged in whorls. Although
the morphology of the stipe extension in P. capitatum is similar to that in
F. multivesiculatum, the conidiomata and conidia are distinct, also
making Falcocladium unsuitable for this fungus.
Because P. capitatum cannot be suitably accommodated in any of
the genera discussed above, we herewith propose a new genus,
Vesicladiella for this fungus.
Vesicladiella Crous & Wingfield gen. nov.
Etym.: Named after the characteristic vesicles resembling those of the
hyphomycete genus Cylindrocladiella.
Coloniae sparsae, albae. Mycelium hyalinum, crassitunicatum, leve,
septatum. Conidiomata sporodochialia, disseminata vel gregaria, alba;
extensiones stipitis erectae, crassitunicatae, irramosae, terminatae vesicis
tenuitunicatis. Conidiophora reducta in cellas conidiogenas. Phialides
Figs 5-13. Conidiophores, conidia and vesicles of Cylindrocladium
naviculatum, Falcocladium multivesiculatum and Vesicladiella capitatum.
Figs 5-8. Cylindrocladium naviculatum. Fig. 5. Conidiophore (bar = 45
pm). Fig. 6. Conidiophore branches with phialides (bar = 9 um). Fig. 7.
One septate conidia (bar = 9m). Fig. 8. Conidiophore branches and a
naviculate vesicle (bar = 9m). Figs 9-11. Falcocladium
multivesiculatum (bars = 9 wm). Fig. 9. Multivesiculate synnematal
conidiophore. Fig. 10. Synnematal conidiophore on a stroma of
chlamydospores. Fig. 11. Falcate conidia with apical and basal
appendages. Figs 12, 13 (bars = 9m). Vesicladiella capitatum. Fig. 12.
Stipe extension with terminal vesicle. Fig. 13. Cylindrical conidia.
455
456
discretae, determinatae, intercalares vel laterales, ampulliformes;
collaricula praesentia. Conidia hyalina, non-septata, levia, cylindrica
extremitatibus rotundatis, guttulata.
Colonies sparse, white. Mycelium hyaline, thick-walled, smooth, septate.
Conidiomata sporodochial, scattered to gregarious, white; stipe
extensions erect, thick-walled, unbranched, terminating in a thin-walled
vesicle. Conidiophores reduced to conidiogenous cells. Phialides discrete,
determinate, intercalary or lateral, ampulliform; collarettes present.
Conidia hyaline, non-septate, smooth, cylindrical with rounded ends,
guttulate.
Holotype: Australia, Queensland, Boombana, M.P., leaf litter of
Araucaria bidwilli, leg. B.C. Sutton & J.L. Alcorn, 27 Aug. 1981, IMI
263277a
Sp. typ. V. capitatum
Vesicladiella capitatum (Sutton) Crous & Wingfield comb. nov. Figs
AO y ES:
Pulvinotrichum capitatum Sutton, Sydowia 41: 338 (1989) (basionym).
Colonies sparse, white. Mycelium hyaline, thick-walled, smooth, septate.
Conidiomata sporodochial, scattered to gregarious, white; stipe
extensions erect, thick-walled, unbranched, up to 160 wm long (from
basal septum to vesicle tip), 2-3 4m wide at the base, terminating in a
thin-walled clavate to ellipsoid vesicle, 3-5 4m wide. Conidiophores
reduced to conidiogenous cells. Phialides discrete, determinate,
intercalary or lateral, ampulliform, 2.5-4 um wide at base, up to 1 wm
wide at apex; collarettes present. Conidia hyaline, non-septate, smooth,
cylindrical with rounded ends, guttulate, 6-9 x 1.5 wm.
The description of two new vesiculate hyphomycete genera from
Eucalyptus and Araucaria leaf litter suggests that there are many genera
on these substrates yet to be discovered and described. Additional
collections and cultural studies with the possibility of new teleomorph
connections would suitably strengthen these arguments, and also provide
Figs 14-19. Falcocladium multivesiculatum (bars = 10 um). Fig. 14.
Multivesiculate synnematal conidiophore. Fig. 15. Penicillate
conidiophore with several stipe extensions. Figs 16, 17. Whorls of
tapering, ampulliform phialides (apices arrowed), forming flacate
conidia. Fig. 18. Falcate conidia with apical and basal appendages. Fig.
19. Conidiophore with falcate conidia arranged in a round packet.
457
458
additional clues as to their possible relationships with other,
morphologically similar genera discussed in this study.
ACKNOWLEDGEMENTS
We wish to thank Dr N.E. El-Gholl (Florida Dept. of Agric & Consumer
Services, USA) and Prof. W.F.O. Marasas (Medical Res. Council,
Tygerberg, South Africa) for their review of the manuscript. We
gratefully acknowledge Drs Brian Sutton (IMI, Bakeham Lane, Egham,
Surrey, TW20 9TY, UK), Bryce Kendrick (Dept. of Botany, Univ. of
Waterloo, Ontario N2L 3G1, Canada) and Walter Gams (CBS, P.O. Box
273, Baarn AG 3740, The Netherlands) for useful advice concerning
Falcocladium, the curator of IMI for loan of material of Pulvinotrichum
capitatum, and Dr Andre Aptroot (CBS) for placing several other
herbarium specimens at our disposal. Dr Keith Seifert (Agric. Canada,
William Saunders Bldg., Ottawa, Ontario, K1A 0C6, Canada) is also
thanked for assistance and advice regarding Falcocladium. We also
acknowledge the South African Foundation for Research Development
for research support to MJW, Miss Carolina de Beer for assistance with
SEM, and Dr Louis van Ryneveld for Latin translations.
REFERENCES
Arambarri, A.M., Gamundi, I.J. & Bucsinszky, A.M. (1981). Microflora
de la hojarasca de Nothofagus dombeyi. II. Darwinia 23, 327-348.
Crous, P.W. (1992). Cylindrocladium and Cylindrocladiella with special
reference to species occurring in South Africa. 318 pp. Ph.D.
dissertation, University of the Orange Free State, Bloemfontein,
South Africa.
Crous, P.W., Phillips, A.J.L. & Wingfield, M.J. (1992). Effects of cultural
conditions on vesicle and conidium morphology in species of
Cylindrocladium and Cylindrocladiella. Mycologia 84, 497-504.
Crous, P.W. & Wingfield, MJ. (1993). A_ re-evaluation of
Cylindrocladiella, and a comparison with allied genera.
Mycological Research 97, 433-448.
Di Cosmo, F., Berch, S. & Kendrick, B. (1983). Cylindrotrichum,
Chaetopsis, and two new genera of Hyphomycetes, Kylindria and
Xenokylindria. Mycologia 75, 949-973.
Rayner, R.W. (1970). A mycological colour chart. CMI and British
Mycological Society. Kew, Surrey, England.
Summerbell, R.C., Seifert, K.A. & Mackinnon, J.A. (1989). Taxonomy
and ecology of Cylindrodendrum album var. paralion vat. nov.
Canadian Journal of Botany 67, 573-578.
Sutton, B.C. (1989). Notes on Deuteromycetes II. Sydowia 41, 330-343.
MYCOTAXON
Volume L, pp. 459-474 January-March 1994
WENTIOMYCES LICHENICOLA
SUBSP. NOV. BOUTEILLEI
CHAMPIGNON LICHENICOLE NON LICHENISE
(DOTHIDEALES, DIMERIACEAE)
par Claude Roux ™*, Olivier Bricaud*,
Emmanuél Sérusiaux ** et Clother Coste ***
*C.N.R.S., U.R.A. 1152, Institut méditerranéen d’écologie et de paléo-
écologie, faculté des sciences et techniques de Saint—Jér6me, F —
13397 MARSEILLE CEDEX 13, France.
**Chercheur qualifié F.N.R.S., Département de botanique, Université
de Liége, Sart—Tilman, B — 4000 LiEcE, Belgique.
***26 rue de Venise, F — 81100 Castres, France.
Résumé
Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp. bouteillei
Bricaud, Roux et Sérusiaux subsp. nov. différe de la sous—espéce liche-
nicola par sa spécificit€ parasitaire (sur Fellhanera bouteillei, excep-
tionnellement sur Bacidina sp.), ses hyphes d’ancrage peu visibles,
moins longues, plus minces et nettement plus claires, et sa répartition
(Europe, Asie et Amérique du Sud). Mise en évidence de périphyses
rudimentaires chez Wentiomyces lichenicola.
Resumo
Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp. bouteillei
Bricaud, Roux et Sérusiaux subsp. nov. diferencas de la subsp. licheni-
cola pro specifa logateco (sur Fellhanera bouteillei, esceptokaze sur
Bacidina sp.), fiksivaj hifoj ne tre videblaj, malpli longaj, pli maldikaj k
distingeble pli helkoloraj, k disvastig0 (Etropo, Azio k S—Ameriko).
Evidentigo de rudimentaj perifizoj Ce Wentiomyces lichenicola.
460
Abstract
Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp. bouteillei
Bricaud, Roux et Sérusiaux subsp. nov. differs from subsp. lichenicola
by parasitic specificity (on Fellhanera bouteillei exceptionnally on
Bacidina sp.), its anchoring hyphae that are hardly visible, shorter, thi-
ner and distinctly paler, and its distribution (Europa, Asia and South
America). Observation of rudimentay periphyses in Wentiomyces
lichenicola.
Introduction
HAWKSWORTH (1980: 385) a décrit trés briévement Wentiomyces liche-
nicola (Hansf.) D. Hawksw., un champignon lichénicole non lichénisé,
connu seulement par les spécimiens originaux récoltés en Ouganda
(Afrique orientale), sur le thalle d’un lichen foliicole non déterminé par
l’auteur. La description de cette espéce a été reprise sans modification
par CLAUZADE et al. (1989).
Dans sa thése sur les lichens épiphytes de Navarre, ETayo (1989)
mentionne un Wentiomyces cf. lichenicola s’établissant sur le thalle de
Fellhanera bouteillei (Desm.) Vézda, rarement sur celui de Parmelia
caperata (L.) Ach. Tout recemment, deux des auteurs du présent travail
(O. BRICAUD ET C. CosTE) trouvaient, dans le Gard et les Pyrénées—
Atlantiques, sur le thalle, ou méme sur les apothécies en mauvais état
de Fellhanera bouteillei, un Wentiomyces quiils attribuaient tout
d’abord, avec C. Roux, 4a Wentiomyces lichenicola, car le matériel récol-
té ne présentait aucune différence avec la description de Wentiomyces
lichenicola donnée par HAWKsworTH Cloc. cit.).
Toutefois, en raison de la briéveté de cette description, l’un de nous
(C. Roux), aidé par O. BRICAUD et E. SERUSIAUX, a fait une étude compa-
rative du matériel type de W. lichenicola UMD et du W. cf. lichenicola
européen. A l’issue de cette étude, il est apparu que le W. cf. lichenico-
la européen —a I|’exclusion du matériel sur Parmelia caperata, qui
n’appartient pas au genre Wentiomyces —differe du W. lichenicola s.
str. par des caractéres peu importants mais constants, si bien que nous
le décrivons ci-aprés comme une nouvelle sous—espéce: W. licheni-
cola subsp. bouteillei Bricaud, Roux et Sérusiaux subsp. nov.
Fig. 1 — Wentiomyces lichenicola subsp. bouteillei (holotype). Périthece
observé dans l'eau, vu de dessus, montrant les cellules polygonales du
péridium (textura angularis), l'ostiole, les poils ramifiés dichotomiquement
et les hyphes végétatives (vers le bas, plus fins que les poils).
461
Diagnose
1 -En latin
Differt a Wentiomyce lichenicola (Hansf.) D. Hawksw. subsp. liche-
nicola hyphis detinentibus perithecia parum conspicuis, brevioribus
(50-100 pm), angustior ibus (2—3):m diametro in basi) et pallidioribus
(pallidofuscis) et quia vivit in Fellhanera bouteillei (Desm.) Vézda
rarissime in Bacidina sp.
Holotypus in herbario C. Roux (MARSSJ n° 21315): Gard, Méjannes—
le—Clap, alt. 300m, in caverna “Trois—Trous”, in thallis Fellhanerae
bouteillei, in foliis Buxi sempervirentis, leg. O. BRICAUD 1992/10/04. Iso-
typus: in herbario O. BRICAUD.
2 —-En langue internationale (espéranto)
Diferencas de Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp.
lichenicola pro periteciaj fikshifoj ne tre videblaj, pli mallongaj
462
(50-100pm), pli maldikaj (¢ebaze 2-3 1m diametraj) kaj pli helkoloraj
(helege ati hele brunaj) kaj pro specifa logateco (€e Fellhanera bou-
teillei, esceptokaze €e Bacidina sp.).
Fig. 2— Wentiomyces lichenicola subsp. bouteillei (holotype). Cellules du
péridium d’aprés un écrasement de périthéce (vue tangentielle).
a) couche externe ; b) couche interne.
463
Holotipo: en herbario C. Roux, MARSSJ: Gard, Méjannes—le—Clap,
alt. 300m, en kaverno Trois—Trous, 22m profonde, sur taloj de Fellba-
nera bouteillei, sur folioj de Buxus sempervirens, kolektis O. BRricAUD
1992/10/04. Izotipo: en herbario O. BRICAUD.
Description
Une description sommaire de Wentiomyces lichenicola subsp. bou-
teillei Bricaud, Roux et Sérusiaux subsp. nov. (sous le nom de W. cf.
lichenicola) a été déja été publiée par Bricaup et al. (1993).
1 -Ascomes (périthéces)
Périthéces (fig. 1) assez petits, de (40)100—200 im de diamétre Cy
compris les poils périthéciaux), isolés, sessiles sur le thalle ou méme
les apothécies en mauvais état de Fellhanera bouteillei, exceptionnelle-
ment sur le thalle d’un Bacidina stérile, noiratres, globuleux, a ostiole
apical, hérissés de poils, ancrés au thalle de l’hdte par des hyphes fixa-
trices non Ou peu visibles au stéréomicroscope mais bien visibles au
microscope a transmission .
Paroi des périthéces (fig. 2 et fig. 3) pseudoparenchymateuse
(= paraplectenchymateuse), a fextura angularis, le plus souvent for-
mée de 2 couches de cellules pigmentées (parfois 3 dans la région
ostiolaire, parfois une seule dans la région basale).
Sur une vue tangentielle (fig. 2), par exemple sur un €crasement de
périthéce, ces cellules apparaissent isodiamétriques ou peu allongées.
La couche externe est formée de cellules (a lumiére de 3-7 x 2—4 pm)
plus petites que celles de l’interne (umiére de 5-10 x 3—7}1m), et les
deux types de cellules présentent une paroi mince (0,5—1 um), pigmen-
tée de brun sombre (pigment dissous dans la paroi), percée de nom-
breux pores (‘Munkporen” des auteurs germaniques).
Sur une coupe transversale (fig. 3), les cellules de la paroi apparais-
sent subrectangulaires (lumiéres de 3-10 x 1—4j1m). Autour de I’ostiole
du périthéce adulte, surtout dans sa partie externe, la paroi des cellules
s’€paissit sensiblement (1-—2,1m) et leur lumiére a tendance a devenir
plus €troite.
La paroi proprement dite, pigmentée, est doublée intérieurement de
2 a 3 couches de cellules incolores, assez laches.
Poils (fig. 4) situés sur la moitié supérieure des péritheces, dressés,
brun sombre, cloisonnés transversalement, de 40-65 ym de longueur,
rigides, ramifiés dichotomiquement deux ou trois fois au sommet, a
paroi €paisse mais amincie au sommet. Dans la zone équatoriale des
464
périthéces, on rencontre parfois des formes de transition entre ces poils
et les hyphes fixatrices (§5).
Ostiole bien visible au microscope (fig. 1 et fig. 3), apical, compre-
nant de l’extérieur vers l’intérieur:
e Une membrane 4 structure pseudoparenchymateuse, qui ferme le
canal ostiolaire avant l’éjection des spores, semblable a celle du reste
de la couche supérieure de la paroi périthéciale mais a cellules plus
petites (umiéres de 2,5—4 x 2—3,5y1m, vue de dessus) et a paroi nette-
ment plus mince (<0,5p1m).
e Un canal ostiolaire, trés court.
e A la partie supérieure de ce dernier, deux couches de cellules en
forme de papilles qui constituent une sorte de couronne (fig. 5). Ce
sont des périphyses rudimentaires, courtes (4-10 x 2—3}1m), tres
Fig. 3— Wentiomyces lichenicola subsp. bouteillei (Gard, Goudargues,
herbier C. Roux, MARSSJ n° 21527). Coupe transversale radiale d'un
périthéce adulte colorée par le bleu de lactophénol, montrant la paroi
(péridium) portant les poils périthéciaux et les hyphes végétatives, pre-
sentant, au sommet de I'ostiole, des cellules modifiێes formant la couron-
ne périostiolaire et contenant des asques a divers stades de leur déve-
loppement ainsi que des restes de paraphysoides (notamment dans la
partie Supéro—axiale).
465
10 um
Fig. 4— Wentiomyces lichenicola subsp. bouteillei (holotype). Poils péri-
théciaux a divers stades de développement.
—
ee
~
~
Fig. 5— Wentiomyces lichenicola subsp. bouteillei (Gard, Goudargues,
herbier C. Roux, MARSSJ n° 21527). Détail partiel des paraphyses rudi-
mentaires garnissant l'ostiole, d'aprés l'examen d'une calotte supérieure
de périthece montée dans l'eau. a) Couche supérieure; b) couche infé-
rieure. En pointillé, contour de l’ostiole.
467
légérementpigmentées (brun jaune pdle ou trés pale), qui présentent
parfois, a leur extrémité distale, une calotte pigmentaire brune. Issues
de la couche externe (sombre) de la paroi périthéciale, elles correspon-
dent aux "périphyses de la formation externe" de JANEx (1971:
439-440), définies antérieurement par CHADEFAUD et AVELLANAS (1967)
et PARGUEY— LEDUC (1967).
A notre connaissance, des périphyses ne semblent pas avoir été men-
tionnées chez les Wentiomyces (en particulier ni par HAwWKsworTH 1980
ni par MULLER et ARx 1962), probablement car elles s’observent difficile-
ment sur une coupe axiale, mais assez bien sur une calotte supérieure
de périthéce vue de l’intérieur ou sur un écrasement modéré d’une
moitié supérieure de périthéce.
De telles périphyses rudimentaires ne sont pas particuliéres au genre
Wentiomyces puisque l'un de nous (C. R.) les a observées également
chez plusieurs espéces de Stigmidium (travail en cours avec D.
TRIEBEL). Elle ont trés vraisemblablement pour réle de régler l’ouverture
et la fermeture de l’ostiole lorsque, a maturité, la membrane supérieure
de l’ostiole a disparu.
2 —-Hamathécium
Constitué par des périphyses rudimentaires (voir ci—dessus) et des
paraphysoides ordinairement peu ou pas visibles car disparaissant pré-
cocement. Chez le périthéce adulte, les restes de paraphysoides ne
sont guére visibles qu’aprés coloration, par exemple par le bleu de lac-
tophénol (fig. 3).
3 - Asques
Asques (fig. 6) claviformes ou subcylindriques, de (35)39-50 x
7-11pm, octosporés, fissituniqués, a sommet €paissi (tholus) et creusé
Fig. 6— Wentiomyces lichenicola subsp. bouteillei. Asques immatures (a)
et matures (b), octosporés (tous observés dans l'eau, le premier en par-
tant de la droite aprés traitement par I’hydroxyde de potassium qui €pais-
sit le tholus et rend la chambre oculaire plus petite mais mieux délimitée).
c) Détail du sommet ascal observé dans le bleu de crésyl (solution
aqueuse a 0,1%), montrant la chambre oculaire et la nasse apicale qui
apparait comme une différenciation de l’endoascus (3 baguettes verti-
cales sur les 4 ont été représentées). a,: holotype; a, et a,:
Pyrénées—Atlantiques, Laruns, herbier C. Coste; b,: Sivergues, herbier
O. BRICAUD ; by_,: holotype; c: Vaucluse, Sivergues, herbier O. BRICAUD.
468
d’une chambre oculaire (surtout visible chez les jeunes asques ou aprés
traitement des asques mirs par K), contenant une nasse apicale (sur-
tout visible aprés coloration par le bleu de crésyl: fig. 6c), 4 paroi I-
(méme aprés pré-traitement par K), 4 ascoplasme I+ (rouge vineux), a
pédoncule trés court ou nul et a pied bilobé, souvent peu nettement,
large.
4-Spores
Spores (fig. 7) par 8, bisériées dans les asques, plus rarement presque
unisériées (herbier Etayo 2939), incolores, de (9,5)10,5— 12, 1-
14,5(16) x (2,5)4-—4, 4-5(6) pm, a rapport longueur sur largeur de
(1,8)2,2—2, 8—3,5(4,0) [n = 58 spores], uniseptées, legerement ou méme
10um
Fig. 7— Wentiomyces lichenicola subsp. bouteille/. Spores vivantes
observées dans le bleu de crésyl (solution aqueuse a 0,1 %). En haut,
Vaucluse, Sivergues (herbier O. BricAupd); torus parfois visible (a droite).
Au milieu, Gard, Méjannes-le-Clap (holotype). En bas, Pyrénées—Atlan-
tiques, Laruns, herbier C. COSTE.
469
nettement rétrécies au niveau de la cloison, parfois aussi legerement
rétrécies au milieu de chaque cellule, contenant 1 ou 2 guttules dans
chaque cellule, dépourvues de halo, mais présentant parfois une calot-
te gélatineuse, trés peu visible, 4 chaque extrémité. Sur le matériel frais,
le bleu de crésyl fait apparaitre un torus équatorial coloré en bleu (fig.
7, en haut et 4 droite) comme la cloison.
Fig. 8— Wentiomyces lichenicola subsp. bouteillei (holotype). Hyphes
d’'ancrage. a) Hyphe relativement courte, entiére, issue d'une cellule du
péridium. b) Extrémité distale d’un autre hyphe, plus long.
Fig. 9— Wentiomyces lichenicola subsp. lichenicola (syntype). Hyphes
d'ancrage (base, milieu et extrémité distale).
470
5 —- Hyphes végétatives
La moitié inférieure des périthéces porte des hyphes d’un brun pale
ou assez clair, d’environ 50—100ym de long et de (1,5)2—3pm de dia-
métre, 4 paroi mince et a cloisons peu nombreuses (fig. 1, 3 et 8).
Cependant, MATZER, qui a également étudié des spécimens de W. liche-
nicola subsp. bouteillei, nous signale (in litt.) que les hyphes fixatrices
peuvent étre incolores et atteignent parfois 130}1m de longueur et 4m
de diamétre.
Les hyphes supérieures, issues de la partie libre des périthéces, sont
essentiellement obliques et horizontales et pénétrent peu profondé-
ment dans le thalle: elles ont probablement un réle essentiellement
fixateur, et on peut les qualifier d’hyphes d’ancrage. Au contraire, les
hyphes issues de la partie la plus basale des péritheéces, en contact avec
le thalle de Il’hdte, s’enfoncent rapidement dans celui—ci (fig. 3) pour
atteindre la couche algale: outre un rdéle fixateur, elles ont vraisembla-
blement une fonction surtout nutritive.
Répartition — Ecologie
Jusqu’ici, nous avons rencontré Wentiomyces lichenicola subsp. bou-
teillei en Espagne (Catalogne, Navarre) et en France méridionale
(Aveyron, Gard, Pyrénées—Atlantiques, Vaucluse). De plus, l’un de
nous (SéRusIAUX) a observé des échantillons d’herbier récoltés dans le
bassin parisien (Val—d’Oise) par BOUTEILLE (voir ci—aprés), mais aucune
récolte récente n’a été effectuée dans cette région. Enfin MATZER (in
litt.) a observé des spécimens de W. lichenicola subsp. bouteillei d’ Asie
et d’Amérique du Sud, dont il précisera la localisation dans une publi-
cation ultérieure.
W. lichenicola subsp. bouteillei se rencontre presque exclusivement
sur le thalle de Fellbanera bouteillei, exceptionnellement sur thalles
stériles d’un Bacidina, lichens sur lesquels il ne provoque aucun dégat
visible, ni macroscopiquement ni microscopiquement: c’est donc,
selon toute vraisemblance un parasymbiote.
Dans le sud de la France, la sous—espéce se rencontre dans d’assez
nombreuses stations de Fellhanera bouteillei, sur feuilles de Buxus
sempervirens, dans le fond de vallons humides soumis a des rosées
abondantes et fréquentes. Dans l’ensemble de son aire européenne, W.
lichenicola subsp. bouteillei se rencontre entre 240 et 600m dialtitude,
aux étages mésoméditerranéen et collinéen, sous un ombroclimat
humide.
471
Affinités
Wentiomyces lichenicola subsp. bouteillei est extreémement voisin de
W. lichenicola subsp. lichenicola, dont elle ne différe que par l’identité
de |’héte, par la longueur, |’€paisseur et la couleur des hyphes fixa-
trices, ainsi que par la répartition géographique (fig. 8 et tableau).
Nous avons étudié dans le détail le matériel type (syntype) de Wen-
tiomyces lichenicola sensu stricto (IMI), qui, rappelons le, provient
d’Ouganda (Afrique orientale). Il est réparti sur 4 feuilles d’Acalypha
fruticosa portant divers champignons foliicoles, la plupart non lichéni-
sés. Wentiomyces lichenicola subsp. lichenicola se rencontre exclusi-
vement sur les thalles du lichen Asterothyrium leucophthalmum (Mull.
Arg.) R. Sant.
Nous avons hésité sur le statut taxonomique de W. lichenicola subsp.
bouteillei: fallait—il le considérer comme une espéce autonome ou
comme une sous—espéce de W. lichenicola?
Quelques arguments plaident en faveur de la premiére hypothése:
l’étude de nombreuses collections du matériel européen nous a montré
que les différences avec le matériel africain, aussi bien morphologiques
(filaments fixateurs) que biologiques (hdte), sont constantes (deux spé-
cimen de la subsp. bouteillei ont cependant été trouvés sur thalles sté-
tiles de Bacidina sp., dans la méme station); en outre, les deux taxons
Ont une aire de répartition bien différente.
subsp. bouteillei
Peu ou pas visibles au
stéréomicroscope
(x 50), non distincte-
ment rayonnantes,
de 50-130 um de long
d'un brun assez incolores, brun pale ou
sombre, brun clair,
de 3-4 um de diametre| de 2-3(4) um de diamé-
dans leur tiers basal | tre dans leur tiers basal
Asterothyrium Fellhanera bouteillei,
leucophthalmum exceptionellement
thalles stériles de
Bacidina sp.
Europe (surtout méri-
dionale), Asie et
Amérique du Sud
subsp. lichenicola
Bien visibles au stéréo-
microscope (x 50), nom-
breuses,
rayonnantes,
Hyphes d’ancrage | de 80-250 um de long,
Afrique orientale
(Ouganda)
Répartition
Tableau: Caractéres distinctifs entre les deux sous-espéces de
Wentiomyces lichenicola.
472.
Toutefois, les caractéres morphologiques séparant les deux taxons
(hyphes d’ancrage plus longues, plus sombres et généralement plus
épaisses chez la sous—espéce type) sont peu importants. Ils résultent
probablement d’une évolution divergente du fait d’un hdéte ou d’un
milieu différent. Par ailleurs, W. lichenicola subsp. lichenicola n'est
connu que de collection type (4 spécimens provenant d’une méme
récolte): la spécificité parasitaire du matériel africain n’est donc pas
certaine.
C’est pourquoi il nous semble préférable de considérer le matériel
européen —et, selon toute vraisemblance, également le matériel asia-
tique et sud—américain— comme une sous—espéce de Wentiomyces
lichenicola: les deux taxons ne different que par des caractéres mor-
phologiques mineurs, par leur spécificité parasitaire (probable mais
non certaine) et par leur répartition geéographique, critéres qui corres-
pondent bien a la notion de sous—espéce. En effet, malgré l’important
matériel de Fellhanera bouteillei examiné en provenance de toutes les
régions du monde par l’un de nous CE. S.) et par MATZzER (in litt), W.
lichenicola subsp. bouteillei n’a pas été observé en Afrique, ou seule la
subsp. lichenicola est connue.
Par ailleurs, HAWKSwoRTH (1980) a décrit une nouvelle espéce, sur le
thalle de Peltigera leucophebia (Nyl.) Gyeln., Wentiomyces peltigerico-
la D. Hawksw., qui différe nettement de W. lichenicola, ainsi que des
autres Wentiomyces non lichénicoles (voir par exemple MULLER et ARX
1962), par ses poils périthéciaux simples.
Spécimens examinés
Note: les dates sont indiquées selon le systéme international :
an/ mois /jour.
1 - Wentiomyces lichenicola (Hansf.) D. Hawksw. subsp.
lichenicola (Syntype)
Uganda, Kiagwe, Mukono, on lichens on leaves of Acalypha frutico-
sa, G. HANsForD 2977 (IMI 5725, syntype !). Spécimen sans date de
récolte (Proc. Linn. Soc. London 157: 186, 1946).
Observations: périthéces de (50)100—150 jm de diamétre (y compris
les poils périthéciaux), munis de périphyses identiques a celle de Wen-
tiomyces lichenicola subsp. bouteillei, asques de 34-47 x 11—13p1m,
spores de 12—15,5 x 3,5-5um (n =10), la plupart avec 2 guttules dans
chaque cellule.
473
2 - Wentiomyces lichenicola subsp. bouteillei Bricaud, Roux et
Sérusiaux
Sauf indication contraire, observé sur thalles de Fellhanera bouteillei
sur feuilles de Buxus sempervirens.
a) Espagne
e Navarre, Aoiz, Poche de Chinchurrinea, U.T.M. 30TXN3344, alt.
550m, 1987/06/12, J. Erayo 2939 (herbier J. ETayo).
e Navarre, Usan, Foz de Arbayun, 30TXN4827, alt. 500m, sur thalles
stériles de Bacidina sp., sur feuilles de Buxus sempervirens,
1991/07/21, J. Erayo 2073 et P. DIEDERICH (herbier J. ETAYO).
e Navarre, Usan, Foz de Arbayun, sur thalle de Bacidina cf. vasakii,
1991/07/12, E. SERUsIAUX, récolt€é avec J. ETayo, P. JAMES et F. ROSE
(LG):
e Catalogne, Girona, Oix, alt. 500m, 1991/11, J. Erayo 5278 (herbier
J. Erayo).
e Catalogne, Girona, Oix, alt. 500m,1991/ 02, P. DIEDERICH Cherbier P.
DIEDERICH).
b) France
e Aveyron, gorges du Lot, carrefour des routes vers Issac et vers
Florentin—la—Capelle, 1986/08, E. SERUSIAUX (LG).
e Gard
— Commune de Méjannes—le—Clap, alt. 300m, dans l’aven des Trois—
Trous, a2 22m de profondeur, 1992/10/04, O. Bricaup [herbier O.
BRICAUD CIsotypus) et herbier C. Roux, MARSSJ n° 21315 (Holotypus)].
— Commune de Goudargue, vallon d’Ussel, alt. 240m, sur thalles et
apothécies de Fellhanera bouteillei, 1992/04/11, C. Roux (herbier C.
Roux, MARSSJ n° 21527).
e Pyrénées—Atlantiques
— Commune de Laruns, Mourat, alt. 600m, 1992/04/01, A. RoYAUD
(herbier C. Coste et herbier C. Roux MARSSJ n° 21287).
— Commune de Sauveterre—de-—Béarn, ile sur le gave d’Oloron,
1985/07, E. SERUSIAUX n° 7508 p. p. (LG).
— Commune de Sainte—Engrace, gorges d’Ujarre, 1991/07, P.
DIEDERICH et J. Erayo (herbier P. DIEDERICH).
e Val—d’Oise (ex Seine—et—Oise p.p.), commune de Magny-en-
Vexin, BOUTEILLE, (UPS, herbier Schaer.).
e Vaucluse, commune de Sivergues, vallée de |’Aiguebrun, alt. 450m,
1992/01/27, O. BricaupD (herbier O. BriCcAuD).
474
3 -“Wentiomyces” sp., sur Parmelia caperata
e Espagne, Catalogne, Gerona, Oix, 500m, sur Parmelia caperata,
1991/11, J. Erayo 11239 (herbier J. Erayo).
Remerciements
Nous remercions tout particuliérement A. BELLEMERE (Paris), P.
DIEDERICH (Luxembourg), J. Erayo (Pamplona) et M. MaTzeR (Graz),
pour l’aide qu’ils nous ont apportée dans cette étude par le prét de spé-
cimens et par leurs commentaires, ainsi que G. CLAUZADE (Cavaillon) et
P. NAVARRO—ROSINES (Barcelona) qui ont relu et corrigé notre manuscrit.
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CHADEFAUD M., 1967.— Remarques sur l’ontogénie et la structure des
périthéces des “Chaetomium’. Le Botaniste, sér. L., Travaux de biolo-
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CLAUZADE G., DIEDERICH P. et Roux C., 1989.— Nelikenigintaj fungoj
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torat inéd., Univ. de Navarra, 980 p.
HawkswortH D. L., 1980.— Notes on some fungi occuring on Peltigera,
with a key to accepted species. Trans. Br. Mycol. Soc. 74(2):
363-386.
JANEX—FAvreE M.—C., 1970 (“1971”). — Recherches sur l’ontogénie,
l’organisation et les asques de quelques pyrénolichens. Rev. Bryol.
lichénol. 37(3): 421.
MULLER E. et Arx J. A. (von), 1962.— Die Gattungen der didymosporen
Pyrenomyceten. Beitrdge zur Kryptogamenflora der Schweiz 11(2):
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PARGUEY—LEDUC A., 1967.— Recherches préliminaires sur l’ontogénie et
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niaux. I. Notions générales. Rev. Mycol. 32(2): 57-68.
MYCOTAXON
Volume L, pp. 475-491 January-March 1994
MISCELLANEOUS NOTES
ON MUCORACEAE
M. A.A. SCHIPPER and R. A. SAMSON
Centraalbureau voor Schimmelcultures ,
P.O.Box 273, 3740 AG BAARN, The Netherlands
ABSTRACT: Three new species of the zygomycete genera Mucor and
Rhizopus are described. M. nanus sp. nov. is characterized by small-sized
sporangia and extremely slowly growing colonies; it resembles some
species of Absidia. M. grandis, sp. nov., resembles M. oblongisporus
Naumov, but differs from it by taller sporangiophores, larger sporangia
and columellae of a slightly different shape and larger sporangiospores. In
addition, it has a higher temperature requirement. Rh. caespitosus, sp.
nov. belongs to the Rh. microsporus complex close to Rh. microsporus
var. microsporus, but is distinct by short sporangiophores in groups of up
to 9 and smooth sporangiospores. Furthermore it can grow at 50°C.
New acquisitions of five strains of M. variosporus, which had only been
known from its type culture from India, are discussed. Zygospores were
obtained and the mating reactions of the strains are given. Mucor faisala-
badensis Mirza et al. takes an intermediate position between Mucor and
Absidia, and the new genus Protomycocladus is proposed. Its apophysate
pyriform sporangia are similar to taxa of the genus Absidia. However,
sporangiophores which arise directly from the substrate, absence of sto-
lons and rhizoids and distinctly stellate zygospores are characteristic of
Mucor.
INTRODUCTION
Over a number of years, some isolates were received which are
considered to be different from the known species of Mucor and
Rhizopus as accepted by Schipper (1973, 1975, 1976, 1978 and
1984) and Schipper and Stalpers (1984). In the present paper
three new taxa are proposed for these isolates. In addition the
type culture of Mucor faisalabadensis was examined and the
taxonomic position is discussed.
476
MATERIAL AND METHODS
Colonies were grown in glass Petri dishes as described by
Schipper (1973) and tall species in 80 mm high dishes to allow
undisturbed development. Unless otherwise indicated, cultures
were grown in the dark. The media used were cornmeal, cherry
decoction, 2% and 4% malt extract, oatmeal, potato-carrot,
potato-dextrose and V8-juice agars.
For the examination and measurements of the various struc-
tures distilled water was used for the preparation of slides. The
branching and the height of the sporangiophores were examined
and measured by a dissecting microscope.
Fig. 1. Mucor nanus (CBS 774.86) a-b. branching pattern of sporangiophores (a.
schematic)
477
Mucor nanus Schipper & Samson, sp. nov. — Figs. 1-2.
Coloniae in agaro maltoso 25°C dilute brunneae, sporangiophora ad 8 ym diam,
irregulariter ramosa; sporangia brunnea, ad 50 ym diam, paries deliquescens et trans-
lucens; columellae subglobosae, ad 21 ym diam, collari minutissimo vel nullo cir-
cumdatae; sporangiosporae ellipsoideae, 1.5-2 x 1 ym; zygosporae ignotae. Crescit et
sporulat temperaturis inter 12 et 30°C.
Typus vivus et exsiccatus: CBS 774.86, isolatus e terra piceeti in Suecia a B.E.
Soderstrom.
Colonies growing extremely slowly, attaining a diameter of up
to 1 cm within 14 days, low, at first white, soon becoming deep
olive-buff. Sporangiophores arising from substrate hyphae,
branching in a mixed fashion, up to 1 mm tall and 8 wm wide.
Sporangia globose, up to 25(-50) wm diam, brownish transpar-
ent, non apophysate, with a globose or slightly applanate-
globose columella up to 21 ym diam, collar absent or minimal;
sporangiospores smooth-walled, hyaline, ellipsoidal 1.5-2 x 1
pm. Zygospores unknown.
Growth and sporulation occurring between 12 and 30°C, opti-
mal between 27-30°C. The various media have no influence on
growth and sporulation at 25°C.
Isolates examined:
CBS 774.86, type culture, and CBS 775.86, both isolated from
coniferous soil in Sweden by B.E. Sdderstrém.
M. nanus is not a typical species of Mucor because of its ex-
tremely slow growth. It is here placed in Mucor on the basis of
the structure of the sporangiophores and sporangia. The species
resembles Absidia zychae Hesseltine & J.J. Ellis, which has
sporangiophores arising either from stolons or from surface or
submerged hyphae and pyriform and globose sporangia. It there-
fore represents an intermediate between Absidia and Mucor. M.
nanus definitely belongs to Mucor, because it differs from
typical species of Absidia by the absence of stolons and by
globose sporangia. The wider temperature range and high
optimum for growth are unusual.
In view of the small size of the sporulating structures, the
name Mucor nanus is proposed. Extremely slow growth in pure
culture of isolates of Zygomycetes may indicate a xerophilic or
mycoparasitic nature (cf. Absidia parricida Renner & Muskat,
478
479
Fig. 2 Mucor nanus, light micrographs. a-c. sporangiophores and sporangia a. x 540,
b. x 660, c. x 620, d. spores, x 1520.
A. verticillata (Beauvérie) Lendner). Cultivation of the type
culture on low water activity media including malt extract agar
with 20 % sugar, showed no improved growth. Furthermore, no
reactions of mycoparasitism were observed in mixed cultures
with other Zygomycetes or with Botrytis cinerea Pers.
Mucor grandis Schipper & Samson, sp. nov. — Figs. 3-4.
Coloniae in agaro maltoso 24°C olivaceo-griseae; sporangiophora 45 pm diam,
primum curvata, deinde recta, plerumque simplicia; sporangia pallide brunnea ad 250
pm diam; columellae conicae vel ellipsoideae, ad 200 x 180 um, collari minutissimo
circumdatae; sporangiosporae ellipsoideae, 12.5-21 x 7.5-12 wm; zygosporae ignotae.
Crescit et sporulat temperaturis inter 21 et 36°C.
Typus vivus et exsiccatus CBS 186.87, isolatus ex excrementis muris a P. C. Misra,
Gorakhpur, India.
Grown on malt extract agar at 24°C reaching the lid (80 mm)
and filling the plate (90 mm) in four days, loosely structured,
clearly showing the substrate hyphae. At first white soon becom-
ing olive buff; reverse yellowish, odour aromatic. Sporangio-
phores rather firm, though occasionally collapsing and produc-
ing marginal colonies, 40-50 um diam, sometimes coiled,
mostly unbranched or infrequently branched, with a few small
droplets occurring on the sporangiophore surface. Young spo-
rangiophores erect, but recurving during development of the
sporangia and straightening again after maturation. Sporangia
pale brownish to yellow, moist to dry, finely echinulate, globose
to slightly applanate, up to 250(-300) um diam in dry condi-
tions, columella conical to cylindrical-ellipsoidal, 115-200 x
100-180 um, with or without brownish contents; collar mainly
consisting of a row of short spines, not very distinct. Sporangio-
spores narrow ellipsoidal with rounded ends, unequal in size,
12.5-21 x 7.5-12 um, smooth-walled, hyaline, with greyish
yellow, cloudy to granular contents.
Temperature requirements: at 17°, in daylight, a bright orange,
but poorly developed colony was produced; between 21 and
36°C good development when grown in darkness; at 40° slight,
480
atypical growth, no sporulation; at 45° no growth.
Mating experiments on malt extract agar at 25 °C:
The type isolate CBS 186.87 was mated with the following
species: Mucor recurvus, CBS 195.71 (+) and Backusella circi-
na, CBS 128.70 (+), which both resulted in incomplete conju-
gations. The mating with Mucor prayagensis, CBS 816.70 (+)
showed only a deep salmon-yellow colouring, which could
indicate a weak reaction. These reactions determine the mating
type of CBS 186.87 as (-). The only available strain of M.
oblongisporus, CBS 569.70, also has a (-) reaction, and so a
possible compatibility between these taxa could not be tested
(see discussion below).
100 pm
Fig. 3. Mucor grandis (CBS 186.87) a. Columellae, b. sporangiospores.
Fig. 4. Mucor grandis, light micrographs. a-b. sporangiophores, x 150, c-d. spo-
rangiospores, x 600.
482
Material examined: CBS 186.87 = P.C.M. 608, ex mouse
dung, Gorakhpur, U.P. India, sent by P.C. Misra, Department
of Botany, University of Gorakhpur, India, in 1977.
M. grandis belongs to the tall Mucor species revised by Schip-
per (1978) and is most closely related to M. oblongisporus
Naumov sensu Naganishi, Hirahara & Joshita (CBS 569.70,
neotype). It differs from M. oblongisporus in the production of
taller sporangiophores, with larger sporangia and columellae of
a Slightly different shape and larger sporangiospores. In addition
it has higher temperature requirements. Recently Chen and
Zheng (1980) also described a species with tall sporangiophores
and spores as M. gigasporus. The type culture (CBS 566.91)
differs from M. grandis by the irregularly shaped sporangio-
spores of up to 30 um, which are borne in relatively small (up
to 90 wm diam) sporangia with globose columellae.
M. grandis and M. nanus can be identified by using the key
given by Schipper (1978: 48-52). M. nanus would fit ad 4:
sporangiospores 1.5-2.0 x 1 um, hyaline and ellipsoidal; M.
grandis ad 46: sporangiospores narrow ellipsoidal, up to 21 wm
in length.
Rhizopus caespitosus Schipper & Samson, sp.nov. — Fig. 5-6.
Coloniae in agaro farina avenae confecto 30°C pallide griseae; sporangiophora 1-9
aggregata, ad 10 ym diam, 200-500 pm longa; sporangia fusco-nigra, ad 60 pm diam;
columellae subglobosae, ad 40 x 35 ym; sporangiosporae angulatae vel breviter ellip-
soideae, leves, ad 7 pm diam; chlamydosporae in mycelio aerio; zygosporae ignotae.
Crescit et sporulat temperaturis inter 15 et 36°C.
Typus vivus et exsiccatus: CBS 427.87, isolatus a M.C. Joshi, Naini Tal, India.
Colonies on oatmeal agar at 30°C attaining a diameter of 9 cm
within 4 days, at first white, soon becoming gray. Rhizoids
simple, subhyaline; sporangiophores on stolons up to 200-500
um in length and 10 um wide, brownish, in groups of 1-9.
Apophysate sporangia brownish black, up to 60(-75) um diam.
Columellae (sub)globose to subglobose-conical, brownish, up to
40 x 35 um, rarely larger. Sporangiospores angular to broadly
ellipsoidal, up to 6-8 um, smooth-walled. Immature long-ellip-
soidal sporangia were observed under all conditions of the
study. Chlamydospores in the aerial mycelium. Zygospores
unknown.
483
‘ ag
b
a
aaa
25 pm
Fig. 5. Rhizopus caespitosus (CBS 427.87). Sporangiophores on stolons and opposite
rhizoids, b. columellae.
Temperature requirements: Growth and sporulation between 15°
and 36 C, slow at 15°C, optimum at 30-36°C. At 45-50°C
growth occurs without sporulation.
Influence of medium: On Czapek agar colonies were thin, with
few sporangiophores.
Material examined: CBS 427.87, isolated from unknown source
by M.C. Joshi, Botany Department (D.S.B. Constituent Col-
lege), Kumaum University, Naini Tal, India.
R. caespitosus has the general characteristics of the R. micros-
porus complex (Schipper & Stalpers, 1984) in which it is near-
est to Rh. microsporus var. microsporus. It differs from this
taxon in the production of sporangiophores in groups of up to 9,
immature long ellipsoidal sporangia which occur under all cul-
ture conditions tested, smooth-walled sporangiospores and the
ability to grow at 50°C. Matings with both mating types of R.
microsporus var. microsporus were unsuccessful.
484
485
Fig. 6. Rhizopus caespitosus a. Sporangiophores x 140, b. sporangiophores with
columellae x 560, c. chlamydospores x 560, d. sporangiospores x 1400.
In the key given by Schipper and Stalpers (1984) the new spe-
cies keys out ad 2: sporangiospores angular to broadly ellipsoi-
dal, up to 8 pm, smooth-walled.
Mucor variosporus Schipper — Fig. 7.
The description of M. variosporus (Schipper, 1978) was based
on a single isolate (CBS 837.70) obtained from rotten fruit of
Ficus. Five strains were later obtained from P.C. Misra
(Gorakhpur, U.P., India): CBS 650.78 = P.C.M. 615 and CBS
651.78 = P.C.M. 616, ex lizard dung, 1977; CBS 652.78 =
P.C.M. 618 and CBS 653.78 = P.C.M. 620, ex shrew dung,
1978; CBS 654.78 = P.C.M. 621, ex owl pellet, 1978.
These isolates differ from the type by producing slightly nar-
rower sporangiospores with more pointed ends. The size of
sporangiospores are as follows:
CBStso 15/0 (1). 4). 0-15.) X 3.0-6: om
CBS 650.78: 7-10 x 2.5-3 wm;
CBS 651.78: 5-10 x 2.5-3 pm;
CBS 652.78: 7-10 x 2.5-3 wm;
CBS 653.78: 5-7 x 2-3 pm;
CBS 654.78: 3.5-7.5 x 2-3 wm
At 36°C all the strains show some growth, but this is rather
slow to restricted.
CBS 650.78 and 651.78 were received as mating partners and
the most profuse yield of zygospores was obtained between
these two. When grown on malt extract agar at 25°C, zygo-
spores were formed throughout the colony at the line of contact.
Zygospores were black, up to 80 um diam (including the
stellate spines 10 um in length) and they were borne between
unequal suspensors. One suspensor is short while the other one
is approximately globose, with or without reddish brown con-
tents.
486
Fig. 7. Zygospore of M. variosporus, x 1400.
Matings of the type strain of the species CBS 837.70 (-) with
CBS 651.78 resulted in the production of similar zygospores and
suspensors, though fewer in number and mixed with some
which were immature and reddish brown. Mating experiments
between all available isolates revealed the following mating
reactions:
(+): CBS 651.78; 652.78;
(-) : CBS 837.70; 650.78; 653.78; 654.78
Mucor faisalabadensis Mirza & al.
Mirza et al. (1979) described M. faisalabadensis based on a
single homothallic isolate and stated that it belongs to Mucor
section Genevensis Hesseltine. Our examination of the type
culture CBS 661.86 showed, that it has sporangiophores which
arise directly from the substrate, and apophysate pyriform
sporangia. The zygospores are distinctly stellate and borne on
unadorned suspensors. The apophysate pyriform sporangia are
487
characteristic of Absidia spp. In common with species of Absidia
subgenus Mycocladus Hesseltine & J.J. Ellis, it has unadorned
suspensors and gradually widening sporangiophores which show
a darker colouring just below the sporangia. In addition, giant
cells in the substrate and temperature requirements for growth
of 15-40(-45)°C are also typical of species of subgenus Myco-
cladus. However, M. faisalabadensis does not fit in Absidia
because the sporangiophores arise directly from the substrate,
while stolons and rhizoids are absent. In addition, the zygo-
spores are not smooth-walled but distinctly stellate as seen in
Mucor spp.
The temperature/growth relations differ from Mucor, where
only few species grow and sporulate well at 40°C and slowly at
15°C, e.g. Mucor indicus, M. prayagensis and M. variabilis.
Since the taxon does not fit either in Mucor or Absidia, the
following new genus is proposed.
PROTOMYCOCLADUS Schipper & Samson, gen. nov.
Sporangia pyriformia, apophysata; sporangiophora simplicia sympodialiter ramosa, ex
hyphis submersis oriunda; stolones et rhizoidea absentia. Zygosporae ornamentatae,
verrucae Mucoris modo stellatae; suspensores aequales haud ornamentati. Species
thermotolerans.
Species typica Mucor faisalabadensis J.H. Mirza, S.M. Khan, Shamsad Begum & S.
Shagufta.
Sporangia pyriform, apophysate, borne on simple sporangio-
phores which are sympodially branched, arising from the sub-
strate; stolons and rhizoids absent; stellately ornamented zygo-
spores borne between unadorned suspensors; thermotolerant.
Type species: Mucor faisalabadensis J.H. Mirza, S.M. Khan,
Shamsad Begum & S. Shagufta
Protomycocladus faisalabadensis (Mirza & al.) Schipper &
Samson, comb. nov. — Fig. 8-9.
Basionym: Mucor faisalabadensis J.H. Mirza, S.M. Khan,
Shamsad Begum & S. Shagufta - Mucorales of Pakistan, Univ.
Agric. Faisalabad Pakistan, p. 69. 1979.
488
100 pm
Fig. 8. Protomycocladus faisalabadensis (CBS 661.86). Young sporangia, b. branch-
ing patterns of sporangiophores, c. columellae, d. sporangiospores, e. zygospores, f.
swollen substrate hyphae.
Fig. 9. Protomycocladus faisalabadensis (CBS 661 .86). a-b. sporangiophores a. xX
670 b. x 600, c. sporangiospores x 1500, d-f. zygospores x 600.
489
490
Colonies grown on malt extract agar at 25°C light brownish
grey; sporangiophores arising from the substrate, unbranched or
with (secondary) branching near the tip due either to restricted
growth conditions (e.g. in a low dish) or to collapse of the
slender stalks, which are approx. 7 wm diam, widening towards
the apophysis, hyaline, darkening below the sporangia; sporan-
gia apophysate pyriform, multispored, up to 75 wm diam,
brownish, with deliquescent walls; columellae cylindrical-ellip-
soidal, up to 50 x 35 wm; sporangiospores ellipsoidal, 3-6(-7.5)
x 2.4-4(-5) um. Zygospores homothallic, reddish brown to dark
brown, mostly inside an exudate droplet, up to 80 ym diam,
including the stellate projections up to 7 wm in length; suspen-
sors opposite, straight, equal, conical, lacking appendages;
substrate hyphae with prominent swellings.
Influence of temperature: between 15-30°C development in-
creasing from very slow at 15°C to luxuriantly at 30°C. At
15°C sporangiospores were up to 4 x 8 um, giant cells occurred
in the substrate; at 33-36°C optimal growth, sporangia and
zygospores present; at 40°C good growth and sporulation, no
zygospores observed, at 45°C a sterile lobed colony was
formed; at 50°C no growth.
Material examined: CBS 661.86, ex type culture (P291) re-
ceived from J.H. Mirza isolated from rodent dung, Faisalabad,
Pakistan.
Acknowledgements
The authors thank Drs Gerald L. Benny and Walter Gams for
reviewing the manuscript. Mrs Karin Vermeulen and Marjan
Vermaas assisted in preparing the illustrations.
REFERENCES
CHEN, G. and ZHENG, R. 1986. A new species of Mucor with giant
spores. Acta mycologica Sinica 1: 56-60.
MIRZA, J.H., KHAN, S.M., BEGUM, S., SHAGUFTA, S. 1979. Muco-
rales of Pakistan. University of Agriculture, Faisalabad, Pakistan.
SCHIPPER, M.A.A. 1973. A study on variability in Mucor hiemalis and
related species. Studies in Mycology, Baarn 4, 1-40.
SCHIPPER, M.A.A. 1975. On Mucor mucedo, Mucor flavus and related
species. Studies in Mycology, Baarn 10, 1-33.
49]
SCHIPPER, M.A.A. 1976. On Mucor circinelloides, Mucor racemosus and
related species. Studies in Mycology, Baarn 12, 1-40.
SCHIPPER, M.A.A. 1978. 1. On certain species of Mucor with a key to all
accepted species. 2. On the genera Rhizomucor and Parasitella. Studies in
Mycology, Baarn 17, 1-70.
SCHIPPER, M.A.A. 1984. A revision of the genus Rhizopus I. The Rh.
stolonifer-group and Rh. oryzae. Studies in Mycology, Baarn 25, 1-19.
SCHIPPER, M.A.A. & STALPERS, J.A. 1984. A revision of the genus
Rhizopus 1. The Rhizopus microsporus-group. Studies in Mycology,
Baarn 25: 19-34.
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495
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496
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497
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AUTHOR'S CHECKLIST
LL] PAGE FORMAT
Y Conforms to one the three formats that are standard for Mycotaxon:
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¥ NO Headers or Page Numbers are printed on manuscript pages.
These may be added after printing using a soft, e.g. , no. 2 pencil. 25
TITLE
Y _Is printed in ALL CAPITAL LETTERS
vY Starts at the prescribed distance from top margin of page 1:
100%- 1.8 cm; Elite- 2cm; Pica-2.4cm
SINGLE-SPACING
Y Is used throughout text of article. At most, a single blank line
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Y No blank lines are used to separate references in literature cited
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ALL ILLUSTRATIONS
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# PHOTOGRAPHS
¥ Photographs after reduction will fill entire width of page.
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SAVE A COPY OF THE MANUSCRIPT FOR YOURSELF
<0 MAIL TO THE EDITOR
Y Cover Letter
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??2?2?2 If you still have questions, contact the Editor-in-Chief, Dr. Jean Boise Cargill,
Mycotaxon, 22 Divinity Ave., Cambridge, MA 02138, U. S. A.
FAX (617) 495-9484 E-Mail cargill@huh.harvard.edu
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499
Author Index, Volume Fifty
Abdullah, N. see Ho, Abdullah and Jalaludin
Albert6, Edgardo and Jorge E. Wright Agaricus pseudoargentinus n. sp. from
Argentina. 271-278
Alfenas, A. C. see Crous, Wingfield, Alfenas and Silveira
Ammirati, Joseph F. see Norvell, Redhead and Ammirati
Archer, Alan W. and John A. Elix Additional new species and new reports of
Pertusaria (Lichenised Ascomycctina) from New Zealand with a revised key to the
corticolous species in New Zealand. 203-217
Ayala, N. see Candusso, Gennari and Ayala
Bagyanarayana, G. and E. John Ravinder A new species of Maravalia from India.
123-125
and A new species of Uromyces on Ocimum from India. 127-129
Bandala, Victor M. see Guzman, Bandala and Montoya
Bhat, R. B. see Thomas and Bhat
Bricaud, Olivier see Roux, Bricaud, Sérusiaux et Coste
Brusse, Franklin A. Four new species in the lichen genus Parmelia (Ascomycotina,
Lecanorales) from Southern Africa, with notes on Southern African lichens. 291-299
Candusso, M., A. Gennari and N. Ayala Agaricales of Baja California - Mexico.
175-189
Cano, J. see Gené, Guarro, Ulfig, Vidal and Cano
Carris, L. M. Observations on Pseudotracylla species. 93-98
Chiarucci, A. and M. G. Mariotti The original collections of Arturo Nannizzi
(1877-1961) in the Herbarium Universitatis Senensis (SIENA). 151-166
Coetzee, J. C. and A. Eicker. Battarreoides diguetti (Gasteromycetes,
Tulostomatales) in Southern Africa. 19-25
Coste, Clother see Roux, Bricaud, Sérusiaux et Coste
Cripps, Cathy and Orson K. Miller, Jr. A new Cortinarius from a mature aspen
stand in Montana. 315-321
Crous, P. W., M. J. Wingfield, A. C. Alfenas and S. F. Silveira.
Cylindrocladium naviculatum sp. nov. and two new vesiculate hyphomycete genera,
Falcocladium and Vesicladiella. 441-458
De Kesel, A. Phaulomyces simplocariae sp. nov. (Ascomycetes, Laboulbeniales) from
Simplocaria semistriata (Coleoptera, Byrrhidae). 191-198
Denchev, Cvetomir M. Validation of the name Microbotryum vinosum
(Ustilaginales). 331
Eicker, A. see Coetzee and Eicker
Elix, John A. see Archer and Elix
Fan, Meizhen and Zengzhi Li. Two new pathogens of dipteran insects. 309-314
A new Cortinarius from a mature aspen stand in Montana.
Gené, J., J. Guarro, K. Ulfig, P. Vidal and J. Cano. Studies on keratinophilic
fungi. I. Chrysosporium pilosum sp. nov. 107-113
Gennari, A. see Candusso, Gennari and Ayala
Giralt, Mireia, Helmut Mayrhofer and Walter Obermayer. The species of the
genus Rinodina (Lichenized Ascomycetes, Physciaceae) containing Pannarin in
Eurasia with a special note on the taxonomy of Rinodina granulans. 47-59
Guarro, Josep see Gené, Guarro, Ulfig, Vidal and Cano
see Pelaez, Polishook, Valldosera and Guarro
Guzman, Gaston, Victor M. Bandala and Leticia Montoya. Tricholosporum
in Mexico and description of a new species. 355-363
, Leticia Montoya, Gerardo Mata and Dulce Salmones Studies in the
genus Pleurotus. III. The varieties of P. ostreatus -complex based on interbreeding
strains and the study of basidiomata obtained in culture. 365-378
Guzman-Davalos, Laura New species of Gymnopilus (Agaricales, Cortinariaceae)
from Mexico. 333-348
500
Hattori, Tsutomu and Leif Ryvarden. Type studies in the Polyporaceae 25.
Species described from Japan by R. Imazeki and A. Yasuda. 27-46
Ho, Y. W., N. Abdullah and S. Jalaludin Orpinomyces intercalaris, a new species
of polycentric anaerobic rumen fungus from cattle. 139-150
Ingleby, K., C. Walker and P. A. Mason Acaulospora excavata sp. nov. - an
endomycorrhizal fungus from Cote d'Ivoire. 99-105
Jalaludin, S. see Ho, Abdullah and Jalaludin
Kamiya, Seigo see Udagawa, Uchiyama and Kamiya
Korf, Richard P. and Pavel Lizon Lambertellinia scutuloides (Sclerotiniaceae), a
new genus and species for a discomycete previously confused with Hymenoscyphus
caudatus. 167-174
Lewis, David P. see Tulloss and Lewis
Li, Zengzhi see Wang, Lu and Li
see Fan and Li
Lizon, Pavel see Korf and Lizon
Lu, Wenhua see Wang, Lu and Li
Lunghini, Dario Menispora convoluta, a new dematiaceous hyphomycete from Central
Italy. 429-433
Mariotti, M. G. see Chiarucci and Mariotti
Mason, P. A. see Ingleby, Walker and Mason
Mata, Gerardo see Guzmén, Montoya, Mata and Salmones
Mayrhofer, Helmut see Giralt, Mayrhofer and Obermayer
Mel'Nik, V. see Nag Raj and Mel'Nik
Miller, Orson K., Jr. see Cripps and Miller
Montoya, Leticia see Guzmdn, Bandala and Montoya
see Guzman, Montoya, Mata and Salmones
Morgan-Jones, Gareth see Phelps and Morgan-Jones
Nag Raj, T. R. Discosia eucalypticola anam.-sp. nov. 349-354
and V. Mel'Nik Redisposals and redescriptions in the Monochaetia-Seiridium,
Pestaltia-Pestalotiopsis complexes. X. Pestalotia granati and Pestalozzina punicae.
435-440
Newberry, Clayton C. see Ririe, St. Clair and Newberry
Norvell, Lorelei L., Scott A. Redhead, Joseph F. Ammirati Omphalina sensu
lato in North America. 1-2. 1. Omphalina wynniae and the genus Chrysomphalina.
2. Omphalina sensu Bigelow. 379-407
Obermayer, Walter see Giralt, Mayrhofer and Obermayer
Pande, Alaka and V. G. Rao Stomiopeltis glochidiicola sp. nov. 199-202
Pelaez, Fernando, Jon D. Polishook, Marti Valldosera and Josep Guarro A
new speciesofDelitschia from West Africa.115-122
Phelps, Ryan A. and Gareth Morgan-Jones. Systematic and biological studies in
the Balansieae and related anamorphs. VIII. TheEphelis anamorph of Balansia
epichloe. 61-72
Polishook, Jon D. see Pelaez, Polishook, Valldosera and Guarro
Pande, Alaka see Pande and Rao
Ravinder, E. John see Bagyanarayana and Ravinder
Redhead, Scott A. see Norvell, Redhead and Ammirati
Ririe, Ted O., Larry L. St. Clair and Clayton C. Newberry. The lichen flora of
Rock Canyon, Utah County, Utah. 323-330
Roux, Claude, Olivier Bricaud, Emmanuél Sérusiaux et Clother Coste
Wentiomyces lichenicola subsp. nov.bouteillei_ champignon lichénicole non lichénisé
(Dothideales, Dimeriaceae). 459-474
Ryvarden, Leif. see Hattori and Ryvarden
St. Clair, Larry see Ririe, St. Clair and Newberry
Salcedo Larralde, I. Contribution to our knowledge of the Aphylloporales
(Basidiomycotina) of the Basque Country. II. 1-7
Salmones, Dulce see Guzman, Montoya, Mata and Salmones
Samson, R. A. see Schipper and Samson
501
Santamaria, Sergio Notes on the genus Monoicomyces (Laboulbeniales,
Ascomycotina): On the species described by Spegazzini. 89-92
Schipper, M.A.A. and R.A.Samson Miscellaneous notes on Mucoraceae. 475-491
Sérusiaux, Emmanuél see Roux, Bricaud, Sérusiaux et Coste
Silveira, S. F. see Crous, Wingfield, Alfenas and Siveira
Simmons, Emory G. Alternaria themes and variations (74-105). 219-270
Alternaria themes and variations (106-111). 409-427
Suarez, V. L. and J. E. Wright Three new Southamerican species of Bovista
(Gasteromycetes). 279-289
Suzuki, Shoji see Udgawa and Suzuki
Thomas, C. M. and R. B. Bhat. Contribution to the lichen flora of Transkei. 9-18
Tulloss,Rodham E. and David P. Lewis Amanita westii - taxonomy and
distribution. A rare species from states bordering the Gulf of Mexico. 131-138
Uchiyama, Shigeru see Udagawa, Uchiyama and Kamiya
Udagawa, Shun-ichi, Shigeru Uchiyama and Seigo Kamiya. A new species of
Ascotricha with non-ostiolate ascomata. 73-80
and Shoji Suzuki. Talaromyces spectabilis, a new species of food-borne
ascomycetes. 81-88
Ulfig, K. see Gené, Guarro, Ulfig, Vidal and Cano
Valldosera, Marti see Pelaez, Polishook, Valldosera and Guarro
Vidal, P. see Gené, Guarro, Ulfig, Vidal and Cano
Walker, C. see Ingleby, Walker and Mason
Wang, Weiman, Wenhua Lu and Zengzhi Li. Furia shandongensis (Zygomycetes:
Entomophthorales), a new pathogen of earwigs. 301-306
Wingfield, M. J. see Crous, Wingfield, Alfenas and Silveira
Wright, Jorge E. see Albert6 and Wright
see Sudrez and Wright
502
INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME FIFTY
This index includes the names of genera, infrageneric taxa, species, and infraspecific
taxa. New names are in boldface, as are the page numbers on which new taxa are
proposed.
Absidia 475, 477, 487
subg. Mycocladus 487
parricida 477
verticillata 479
zychae 477
Abstoma 289
Acanthobasidium
phragmitis 1-4
Acarospora
boulderensis 323, 326
glaucocarpa 326
nodulosa
var. nodulosa 324, 330
strigata 326
Acaulospora 99, 103-104
cavernata 103-104
excavata 99-100, 102-103
foveata 103
lacunosa 103
paulinae 103
scrobiculata 103-104
undulata 103
Agaricus 271, 277-278, 404
sect. Campestres 276
sect. Clitocybe 176
sect. Sanguinolenti 276
argentinus 275-276
bisporus 404
chrysoleucus 388-389
ectypoides 396
fuscofibrillosus 277
grossulus 380-381, 388
hoffmanii 390
impudicus 276
lanipes 276
melizeus 389
pinophilus 176
pseudoargentinus 271-272, 273, 275-
276
umbelliferus 388-389
var. abiegnus 380, 388
xanthophyllus 392
Alternaria 163, 219-223, 226-232, 234,
238, 240-244, 246-248, 250,
292-253; 259, 298, 200, 262;
264, 266-270, 409-411, 413,
422, 426-427
acalyphae 219, 239-240, 260, 267
acalyphicola 219, 260-261, 267
agripestis 219, 255-257, 266-267
alternata 220, 413, 422
angustiovoidea 246, 255-256, 266-267
cheiranthi 241
cicina 219, 264-265, 267
compacta 223, 230, 237
conjuncta 410
constricta 219, 252-253, 254, 267
crotonis 241
dianthi 241
ethzedia 410
euphorbiae 220, 225-226, 228-229,
242-245, 247-250, 252, 267
euphorbiicola 220, 231, 244-246,
248, 267
heveae 219, 262-263
infectoria 409-411, 413, 415, 417-
418, 420, 422, 424
metachromatica 409, 418, 420
mouchaccae 264
obtecta 219, 229, 250-252, 267
oregonensis 409, 417-418
ricini 233-235, 237-238, 241, 266-267
subulata 219, 258-259, 267
tenuissima 241
triticicola 409-410, 424, 426
triticimaculans 409, 413, 415, 417-
418
triticina 409-410, 422, 424, 426
zinniae 243
Amanita 131, 137-138
sect. Lepidella 138
sect. Validae 137
flavorubescens 137-138
pachysperma 138
rubescens 133
sculpta 137
subvirginiana 138
virginiana 138
westii 131-134, 136-138
Amaurascopsis 113
{[Amaurascopsis] perforatus 112-113
Anaeromyces 139, 150
elegans 149
mucronatus 150
Anthostomella
maritima 155
Antrodiella 35
gypsea 27, 32, 35
Aporomyces 192, 198
Arrhenia 398, 406
Aschochytella 156
deformis 156
vaginarum 155
Ascotricha 73-74, 76, 78
amphitricha 74
distans 73-74, 76-77, 79
erinacea 77
Ascotrichella 74
hawksworthii 74
Aspicilia
calcarea 326
cinerea 326
desertorum 326
Asproinocybe 362
Asterothyrium
leucophthalmum 471
Atkinsonella
hypoxylon 61-62, 64, 72
Auriporia
aurulenta 1-4
Australohydnum
dregeanum 41
Bacidina 459-463, 470-471, 473
vasakii 473
Backusella
circina 480
Balansia 61-62, 64, 70, 72
aristidae 61, 64, 70, 72
claviceps 64, 72
cyperi 61, 72
epichloe 61-62, 64-66, 68-70, 72
henningsiana 62, 70
obtecta 64
strangulans 64
Batarrea
digueti 19
stevenii 19-20, 22, 24
Battarrea 20-22
digueti 24
f. minor 20
griffithsii 20
Battarraeastrum
503
digueti 20
Battarreoides 19, 21-22, 24
diguetii 19-21, 24
potosinus 20, 24
Bolbitius
vitellinus
var. fragilis 175
Botrydina 405
Botryosporium 222
diffusum 222
longibrachiatum 222
prorumpens 220-222
Botrytis
cinerea 479
Bovista 24, 279-280, 284, 287-288
subg. Bovista 287
subg. Globaria 280
sect. Nannobovista 287
ser. Fuscae 287
ser. Polymorphae 280
aenea 281
aestivalis 288
var. perverrucispora 280
coprophila 288
cunninghami 281-282
delicata 284
dermoxantha 284
fuegiana 279, 281, 286
fulva 287
glacialis 284
oblongispora 284, 288-289
polymorpha 280
pusilla 288
singeri 279, 282, 284, 286
sublaevispora 279, 284, 286, 288
Buellia 59
Bulgaria 172
Bulgariella 172
Bullatina 17
Byonora 18
Byssochlamys 81
Calcarisporium 78
Calicium
salicinum 55
Caloplaca
cerina 55
citrina 326
holocarpa 326
trachyphylla 326
Calvatia
versispora 29
Camarophyllus 379, 388-389, 404
504
[Camarophyllus] subg. Aeruginospora
389
grossulus 380-381
pratensis 389
Camarosporium
gnidii 156
obtusum 156
psammae 156
Candelariella
deflexa 326
rosulans 326
vitellina 55
Cantharellula 396
Cantherellus 390, 396
Cantharomyces
bordei 192
Catapyrenium
granulosum 326
lachneum 326
Catinella 172
Cattaena 268
Chaetomium 76, 78, 474
Chaetopsis 458
Chrysomphalina 379-380, 389-390,
394, 396, 398, 400-401, 405-
406
aurantiaca 390-392, 398
chrysophylla 385, 390, 392, 396, 401
var. chrysophylla 390-392, 398
var. hoffmanii 379, 390-392, 398
var. salmonispora 379, 390-392,
398
grossula 379-380, 381, 383-392, 398
strombodes 390, 392
Chrysosporium 112-113
merdarium 112
pilosum 107-108, 110, 112-113
vallenarense 112-113
Cladonia
subpityrea 208
Cladosporium
anonae 163
Clathrospora 154, 165
patriniae 154
Clitocybe 175, 187-188, 362, 379, 394,
399, 401, 404-406
sect. Candicantes 383
acerba 178
americana 178
californiensis 178
fragrans 176
pinophila 175, 177-178, 189
subbulbipes 178
trullaeformis 178
venustissima 404
xanthophylla 388-389, 392
Codinaea 433
Collema
coccophorum 327
flaccidum 327
polycarpon 327
Coltricia 45
cinnamomea 43
dependens 43
pusilla 43
Coltriciella
pusilla 43
Comoclathris 165
Coniochaeta 74
Coniothyrium 163
garriae 156
garryae 156
manihotis 156
Coprinus
lagopides 178
Coriolellus
kusanoi 37
Corticium
dregeanum 41, 43
Cortinarius 315-317
subsect. Multiformes 318
stirps Multiformis 318
stirps Rapaceus 318
albidus 318
aleuriosmus 317
amarescens 318
hedyaromaticus 315-316, 318-320
lustratus 317
multiformis 318
ochropallidus 318
osmophorus 317
rapaceus 318
talus 318
Coryneum
feijoae 157
Crocicreas 172-173
Cryptoderma
cercidiphyllum 31
citrinum 31
yamamoi 44
Cryptosporium 157
lunatum 157
Cryptostegia 124
Ctenomyces
trichophyticum 166
Curvularia 223
Cyathicula 172-173
hyalina 172
505
Cylindrocladiella 441-442, 444, 447, funiculosa 78
454, 458 pulvinata 78
camelliae 452 Didymochaeta
elegans 452 atropae 157
Cylindrocladium 441-444, 446-447, Didymosphaeria
458 brunneola 158
avesiculatum 442, 446 Dimelaena 59
candelabrum 442-443, 446 oreina 327
clavatum 442, 446 thysanota 323, 326-327
colhounii Diplodia
var. colhounii 452 dorycnea 157
gracile 442, 446 portlieriae 157-158
hawksworthii 442, 446 smilacella 158
naviculatum 441, 443-444, 446, 454 Diplodina
pteridis 442, 446 odontitidis 158
scoparium 442, 446 odontitis 158
Cylindrodendrum 441-442, 444, 447, vaginarum 155
452, 454 Diplotomma
album 452 alboatrum 327
var. paralion 452, 458 Dirinaria 48-49, 56
Cylindrotrichum 458 picata 12, 14
Cyphelium Discosia 352, 354
tigillare 55 arxii 354
ceanothi 353
eucalypti 353
Daedalea 33 eucalypticola 349-350, 353
dickensii 30, 33, 35 lauricola 353
incana 33, 35 novaezelandiae 353
kusanoi 37 subramanianii 354
quercina 35 Dothichloe 64
Daedaleopsis
conchiformis 30-31
confragosa 33 Echinodontium 45
nipponica 39 ballouii 36
purpurea 39 japonicum 34, 36-37
styracina 39 Embellisia 220, 242, 269
tricolor 33, 39 conoidea 242
Datronia Entoloma
stereoides 43 lividoalbum 179
Delitschia 115-116, 120-122 sericeum
confertaspora 116, 118-121 var. sericeum 179
dochmiophragmia 120-121 Entomophthora 314
ionthada 121 aphidis 314
mesostenospora 121 aquatica 313
myriaspora 121 brevinucleata 314
polyspora 121 crustosa 306
sexdecimspora 121 forficular 304-305
spiralirima 120-121 Ephelis 61-62, 64-66, 68-70, 72
Dermatocarpon borealis 64
intestiniforme 327 japonica 64
miniatum 327 mexicana 64
reticulatum 327 oryzae 64
Dictyosporium 268 trinitensis 64
Dicyma 73-75, 78 Epicoccum 222
506
Eriosphaeria
dumetorum 155
littoralis 155
Erynia 304-306, 308, 313-314
aquatica 313
chironomis 307, 311-313
conica 312-313
ellisiana 304-305
forficulae 301, 305
var. major 304
ithacensis 306
pieris 306, 314
rhizospora 312-313
virescens 306, 314
Eryniopsis 314
Erysiphe
polygoni 164
f. robiniae-hispidae 155
Eubalansia 64
Eupenicillium 81
Falcocladium 441, 447, 452, 454, 458
multivesiculatum 448, 450, 454, 456
Fellhanera
bouteille: 459-463, 470-473
Flavoparmelia 18
Flaviporus 6
brownei 1, 4
Fomitopsis
castanea 29
kiyosumiensis 37
Fulgensia
fulgens 327
Furia 304-305
crustosa 305
ellisiana 301, 304
ithacensis 305
shandongensis 301-302, 303, 305,
313-314
Gaetneriomyces 149
Galerina 337
Ganoderma
colossus 44
lucidum 39, 43
neo-japonicum 39
subumbrachulum 43
tsunodae 42, 44
Geniculosporium 76
Gerronema 188, 379, 389, 392, 394-
397, 400-401, 406
sect. Fibulae 397
sect. Haasiella 396
grossulum 380
marchantiae 398
melanomphax 395, 401
strombodes 379, 388, 392-393
subclavatum 398
xanthophyllum 379, 392-393
Glenosporella 165
Glomus
constrictus 104
Gonohymenia
nigritella 327
Graphyllium 165
Grifola 45
albicans 29
frondosa 29
Gymnoascus
gypseus 164
Gymnopilus 333, 348
aurantiacus 340
aurantiophyllus 340
commune 333-335, 336
galerinopsis 333-334, 336-337
hemipenetrans 333-334, 338-339,
340, 347
liquiritiae 336
longipes 336
magnificus 343-344
magnus 343-344
medius 333, 338, 340, 343
oregonensis 337
penetrans 340
picreus 337
radicicola 347
robustus 333, 338, 342-343, 344
rugulosus 344
sapineus 340
spectabilis 343-344
stabilis 340
subbellulus 347
subsapineus 336, 340
subspectabilis 344
tuxtlense 342, 345-347
Haasiella 379, 394, 396, 400, 402,
405-406
splendidissima 396, 405
venustissima 396, 402
Hamigera 81
Hansfordia 78
pulvinata 78
Hebeloma 316
Helminthosporium
{Helminthosporium] allii 269
Helotium 170
berberidis 168, 171
Hemimycena 397
Hendersonia 158
arenicola 158
cyperi-aegyptiaci 158
pteleae 158
Hendersonula 158
Heterobotrys 164
Heterodermia 12
diademata 12, 14
Heterosporium 234
Hohenbuehelia 371
Humicola 74
Hydnochaete 46
tabacinoides 44
Hygrocybe 379, 399
sect. Firmae 405
luteo-omphaloides 379, 399-400
occidentalis 399
parvula 399
Hygrophorus 388, 399, 404
sect. Hygrocybe 399
mycenoides 399
occidentalis 399
wynniae 380-381, 388
Hymenoscyphus 167, 170-174
caudatus 167-168, 170-171 173-174
scutulus 170
subserotinus 172
Hyperphyscia
adglutinata 327
Hypocenomyce
scalaris 55
Idriella 167, 170, 172-174
Inonotus 46
flavidus 43
sciurinus 43
Irpex 46
parvulus 41
purpureus 41
tabacinoides 44
Karoowia 292, 298
Keratinomyces
ajelloi 164
Kochiomyces 149
Kylindria 458
507
Laboulbenia
slackensis 198
Laccaria 188
bicolor 179
Lactarius 181, 187
alnicola 175
var. alnicola 179-180
var. pitkinensis 181
var. pungens 181
argillaceifolius
var. megacarpus 175, 180-181
olympianus 181
payettensis 181
psammicola
f. glaber 181
yazoonensis 181
zonarius 181
Laetiporus
versisporus 29
Lambertella 170-172
Lambertellinia 167-168, 170-172
scutuloides 167-168, 169-172
Lanzia
serotina 172, 174
Lecanora 48, 51, 54, 59
dispersa 327
hageni 327
hueiana 50
muralis 327
symmicta 55
thallophila 327
varia 55
Lecidea
leucothallina 327
turgidula 327
Lecidella
stigmatea 327
Lentinus
sajorcaju 372
Lenzites
tricolor 33
vespacea 37
Lepiota
americana 133
cristata 276
Lepista 359
sordida 359
Lepraria
incana 327
Leptoglossum 398, 406
Leptoporus
mollis 36
Leptosphaeria 154
altaica 154
508
Lewia 410
Lycoperdon 282, 284
Macrolepiota
rachodes
var. bohemica 182
Macrophoma 159, 166
leptopoda 159
Macrospora 165
Macrosporium 163, 220, 223, 248
acalyphae 219, 238-240
cavarae 230
commune
f. Ricinis communis 224
compactum 222-223, 230, 237
euphorbiae 219, 224-231, 242-250,
252-253
phoenicis 163
ricini 231-234, 237, 270
rosarium 234
var. piscariae 232
trichellum 224
var. caulicola 223-224
var. "caulicolum" 223
truncatum 243
Malbranchea 113
filamentosa 112
Marasmiellus 397
sect. Fibulae 397
Maravalia 123-125
cryptostegiae 124
ramacharii 123-124
Megaspora
verrucosa 327
Meionomyces 197
Melanielia
exasperatula 55
olivacea 55
Melanoleuca 188
Melanoporia
castanea 27-29, 31
nigra 31, 45
Menispora 429, 432-433
britannica 432
caesia 432
convoluta 429, 431-432
gamsii 432
uncinata 432
Menisporopsis 433
Microbotryum
vinosum 331
Microdochium 173
Micromphale
inodorum 182
Micropsalliota 277
Microsphaeropsis 161
Microsporum
gypseum 164-165
nanum 164
Milvina 48, 54-55
Monochaetia 164-165, 435
Monoicomyces 89
affinis 89
britannicus 90
caloderae 89
homalotae 89-90
infuscatus 89
invisibilis 89
ocaleae 89
ternatus 89-90
unilateralis 89-90
venetus 89-90
Morularia 164
Mucor 475, 477, 482, 487, 490
sect. Genevensis 486
circinelloides 491
faisalabadensis 475, 486-487
flavus 490
gigasporus 482
grandis 475, 479-480, 482
hiemalis 490
indicus 487
mucedo 490
nanus 475-477, 479, 482
oblongisporus 475, 480, 482
prayagensis 480, 487
racemosus 491
recurvus 480
variabilis 487
variosporus 475, 485-486
Mycena 397, 399, 407
Mycocladus 487
Mycoleptodiscus 98
Mycosphaerella 154
Nannizzia
incurvata 165
Neocallimastix 139, 147, 149-150
joyonii 139, 141, 150
variabilis 149-150
Neosartorya 81
Neozygites 314
Nesolechia 296
Nodulisporium 76
Nothopanus
lignatilis 183
Numbya 229-230, 248
Oidium 164
monosporum 155
orbiculare 155, 164
Omphaliaster 362
Omphalia 187-188, 385, 405-406
abiegna 380-381, 383, 385, 388
bibula 380, 385
var. citricolor 380-381, 388
f. citricolor 380
bresadole 388, 392
chrysophylla 405
hohensis 398
hypoxantha 388, 392
occidentalis 399
umbellifera 385
var. bresadolae 388
var. chrysoleuca 388
wynniae 380-381, 385, 388
xanthophylla 392
Omphalina 175, 184, 187-188, 379,
388-389, 394-400, 402, 404-406
abiegna 380-381, 388
bakeri 184
bibula 380, 388
californiensis 184
chrysophylla
var. chrysophylla 398
var. salmonispora 390, 398
epichysium 397, 406
ericetorum 184, 385, 398
grossula 380-381, 389
hoffmanii 398
hohensis 379-398
hudsoniana 398
hydrogramma 397
illudens 186
jalapensis 184
luteicolor 398
luteovitellina 398
Mac Murphyi 184
marchantiae 379, 398
niveicolor 184
occidentalis 379, 399
olearius 186
olivaria 399
olivascens 186
var. indigo 184
postii 398
pyxidata 397-398, 402, 406
rosella 175, 183-185, 189
sphagnicola 398
509
sphagnophila 398
subclavata 184, 398
tepeitensis 184
umbellifera 184, 397
var. citrina 380-381, 388
viridis 398
wynniae 379-381, 388-389, 398
Omphalotus 187
Onychocola
canadensis 112-113
Orpinomyces 139, 141
bovis 139, 141, 149-150
intercalaris 139-141, 148-149
joyonii 141, 149-150
Oxyporus
cuneatus 35
Pachysporaria 48-49, 56
Paecilomyces 81, 87-88
spectabilis 82
variotii 88
Pandora 308, 310
chironomis 310
shaanxiensis 307-308, 309-310, 313
Parasitella 491
Parmelia 12, 18, 291, 295-298
subg. Flavoparmelia 12
subg. Parmotrema 12, 14
subg. Punctelia 12
subg. Rimelia 12
subg. Xanthoparmelia 12, 14, 295, 298
adhaerens 292, 295
austrosinensis 12, 14
borreri 12
brevilobata 297
caperata 460, 474
chionophila 291-293, 298
emolumenta 291-292, 293, 298
endochromatica 296
endomiltodes 297
evernica 296
fissurina 297
lecanoracea 297
mesmerizans 291, 294, 298
molybdiza 296
mutabilis 12, 14
parilis 296-297
ponderosa 296
protoquintaria 291, 294, 297
reticulata 12
rubropustulata 297
saniensis 297
saxatilis 50
510
[Parmelia] soredians 12
subcrustacea 292, 297
sulcata 55
supposita 293-294, 295, 298
tantillum 294
xanthomelaena 294
Peltigera 59, 474
canina 327
leucophlebia 472
Penicillium 88
Pentopetia 124
Perenniporia 39
fulviseda 36
japonica 27, 36
latissima 37
minutissima 27, 37-38, 43
ochroleuca 39
Pertusaria 203-206, 215, 217
albissima 203, 207, 210, 216
alboatra 203, 211, 216
barlettii 203-205, 217
celata 203-204, 205-206, 217
circumcincta 216
coronata 213
cretacea 210-211
erumpescens 206
globulifera
ssp. glaucomopsis 209
* glaucomopsis 209
hadrospora 203, 205-206, 216
laevis 210, 216
leucodeoides 210-211
leucodes 210-211, 217
var. inconspicua 210
leucoplaca 203, 211-212, 216
leucoplacoides
var. major 306
leucostigma 212
melaleuca
var. heterochroa 213
melaleucoides 207, 211, 216
nigrodisca 209
novaezelandiae 209, 215-216
parvula 203, 205-206, 207, 216
psoromica 203, 205, 207-208, 215
scaberula 208
schizostomella 203, 212-213, 216
scutellifera 203, 205, 208-209, 216
sorodes 206, 216
sphaerophora 209
sphaerulifera 208
sporellula 203, 205, 209, 216
subisidiosa 203, 213, 216
subplanaica 203, 214, 216
tetrathalamia
var. major 206
thamnolica 203, 214-216
theochroa 210-211, 217
truncata 209, 215-216
velata 203, 215-216
Pestalopezia 172
Pestalotia 159, 164-165, 435
granati 435-436
jodhpurensis 438
ventricosa 159
Pestalotiopsis 159, 161, 165, 435-436,
438-439
granati 436-439
Pestalozzina
punicae 435, 438
Phaeociboria 172
Phaeohelotium 172
Phaeophyscia
ciliata 328
endococcina 328
nigricans 328
orbicularis 328
Phaeoseptoria
rubiae 159
Phaeotellus 395, 398
Phaulomyces 191-192, 197
denticulatus 191
octotemni 191, 197
perparvus 191
simplocariae 191-192, 193, 195-197
Phellinus
conchatus 31
kanehirae 37
nilgheriensis 37
pini 44-45
vorax 44
xeranticus 31
Phlegmacium 315, 318, 321
subsect. Multiformes 318
amarescens 318
talus 318
Phoma 160, 166
debeauxi
f. scapicola 159
fimbriata 159
herbarum
f. loti-cretici 160
schini-molli 160
Phomatospora
arenaria 174
berkeleyi 174
Phomopsis:
phellodendri 160
Phyllosticta 160, 166
aberiae 160, 164
altaica 160
helwingiae 160-161
ibotae 161
Phyllostictella 161
dracaenae 161
draconis 161
Physcia 48-49, 51, 53-57
caesia 328
dimidiata 323, 326, 328
dubia 328
opuntiella 59
stellaris 14
tenella 328
Physconia 55
detersa 328
grisea 328
muscigena 328
Phytoconis 379, 394, 397-398, 400,
402, 405-406
botryoides 403
ericetorum 385, 387, 398, 403
luteovitellina 389, 398
viridis 398
Piptoporus
soloniensis 37
Piromyces 147, 150
communis 149-150
spiralis 150
Pithomyces 223, 230, 237, 242
chartarum 223, 237
Platysporoides 165
patriniae 154
Pleospora 165
comata 154
herbarum 159
f. rosae-banksiae 154-155
Pleurotus 188, 365, 367, 375-377, 405
columbinus 367, 370-371, 373-375
cornucopiae 375
cornucopioides 375
cystidiosus 377
djamour 371, 377
"florida" 371, 374
floridanus 371, 374
ostreatus 365, 367-368, 370-377
var. columbinus 365-368, 371, 374
var. florida 365, 367, 371, 374
var. ostreatus 365-367, 370-372,
374-375
pulmonarius 365, 367, 371-376
sajorcaju 365, 367, 372, 374
smithii 374, 377
Pluteus
nanus 186
Poculum 172
Podocrea 24
Polycoccum 296
Polydiscidium 172
Polyporus 29, 46
calvatioides 29
flavidus 43
kanehirae 37
komatsuzakii 37
nigra 31
nilgheriensis 37
sendaiensis 43
tsunodae 44
versisporus 29
Polystictus
gypseus 35
nipponicus 39
orientalis 41
Poria
fulviseda 36
Porina 17
Protodaedalea
hispida 35
Protomycocladus 475, 487
faisalabadensis 487-488
Psalliota 278
ey
Pseudoarmillariella 379, 394, 396, 399,
403
ectypoides 396, 403
Pseudomicrodochium
candidum 452
Pseudotrachylla 93-94, 97-98
dentata 93-95, 97-98
falcata 93-95, 97-98
Psilocybe
coprophila 186
Psiloparmelia 298
arhizinosa 296
Psora
tuckermanii 328
Puccinia 128
Pulveria 80
Pulvinotrichum 442, 444, 447, 452
album 452
capitatum 452, 454, 456, 458
Pyrroderma 45
sendaiense 43
Ramalina 16
aspera 14, 16
celastri 14, 16
at
Renispora
flavissima 112-113
Rhabdospora
rubiae
var. amerospora 161
Rhizomucor 491
Rhizopus 475, 491
caespitosus 475, 482-483, 485
microsporus 475, 483, 491
var. microsporus 475, 483
oryzae 491
stolonifer 491
Rickenella 379, 394, 397, 399, 403,
405-406
fibula 397, 403, 405
swartzii 397
Rimbachia 406
Rinodina 47, 49-51, 55-56, 58-59
adirondackii 48, 56
archaea 52
beccariana 56, 59
confragosa 56
dalmatica 47-49, 56
efflorescens 47, 49-50
excrescens 47-53, 58
granulans 47-49, 52, 54-55, 58
granuligera 48, 56-57
hueiana 50
marysvillensis 48, 56-57
var. thujae 57
murrayil 48, 56-57
poeltiana 58
pruinella 47-49, 55-57
roboris 56
santorinensis 47-48, 56
sibirica
var. granulans 54
sp. 1 50
thujae 48, 51-52, 57
Rinodinella 59
Rhizocarpon
geminatum 328
Rhizoplaca
melanophthalma 328
peltata 328
Rinodina
bischoffii 328
pyrina 328
turfacea 323, 326, 328
Robillarda 161
aquatica 161
Ruminomyces 150
elegans 149-150
Russula
cyanoxantha 182
Rutstroemia 172-173
Sarcodontia
crocea 1, 4, 6
setosa 6
Sclerocystis
clavispora 104
Scutellospora
castanea 105
Seimatosporium 157
Seiridium 435
Septoria 160
asteris-alpinis 161-162
iridis-japonicae 162
moesiaca 162
oligocarpa 162, 164
pachypleuni 162
Seuratia 164
Sorokina 172
Speira 268
Sphaerella 154
agostinii 153
pachypleuri 162
sassafras
f. major 154
Sphaeriopsis 159
Spizellomyces 149
Sporothnix 174
Stagonospora 158
schoeni 162
Steccherinum 41, 46
Stemphylium 220-221, 223-224, 240,
243, 248, 266
botryosum 240
var. majus 240
floridanum 241
var. euphorbiae 240
lycopersici 240-241
pyriforme 223
solani 243
trichellum 223, 268
Stigmidium 467
Stomiopeltis 199, 201-202
aspersa 199, 202
glochidiicola 199-202
polyloculata 199, 201-202
Strobilomyces 137
Talaromyces 81, 87-88
sect. Emersonii 81
byssochlamydoides 87-88
{Talaromyces] leycettanus 87-88
spectabilis 81-82, 83-88
Tarichium 314
Teloschistes 17, 59
exelis 14
hypoglaucus 14
Thomophagus 44
Toninia
caeruleonigrecans 329
Torula
altaica 163
Trachyderma 44
Trametes 39
dickinsii 33
japonica 36
kusanoana 37
kusanoi 37
lactinea 41
minutissima 37
orientalis 41
sendaiensis 43
symploci 43
tricolor 31, 33
Trichaptum 41
byssogenus 41
fuscoviolaceus 41
parvulus 27, 40-41
Tricholoma 355, 358, 360, 362-363
subg. Tricholosporum 355
columbretta 186
porphyrophyllum 359
Tricholosporum 355, 359-360, 362
atroviolaceum 358
longicystidiosum 355, 361-362
porphyrophyllum 355, 359-361
pseudosordidum 355, 358-361
subporphyrophyllum 355, 359-360
tropicalis 355-356, 357, 360
Trichophyton
gypseus
var. asteroides 166
terrestre 164
Triparticalar 149
Trochila 172
Trogia 395
Truncatella 159, 165
Tubakia 97
dryina 98
Tyromyces 36
incarnatus 34-35
Ulocladium 220, 223, 242
manihoticola 241
513
Umbilicaria
phaea 329
torrefacta 329
virginis 329
Uncigera 442, 447, 452
Uredo
vinosa 331
Uromyces 127-128
lami 128
ocimi 128
orthosiphoniis 128-129
prunellae 128
ramacharii 127-129
valerianae 128
Usnea
undulata 14, 16
Ustilago
vinosa 331
Velutarina 172
Venenarius
westii 133
Verrucaria
muralis 329
Verticicladium 78
Vesicladiella 441, 454
capitatum 452, 454, 456
Volvariella 187
gloiocephala 186
Wentiomyces 460, 467, 472, 474
lichenicola 459-461, 463, 471-472
ssp. bouteillei 459-460, 462-465,
467-473
ssp. lichenicola 459-461, 469, 471-
472
peltigericola 472
Xantholinus
gracilis 89
Xanthoparmelia 298
aerolata 296
coneruptens 296
eruptens 296
evernica 296
inconspicua 294
plittii 329
protoquintaria 297
pustulifera 296
pustulosorediata 296
rubropustulata 296
514
[Xanthoparmelia] saiensis 296
xanthomelanoides 293
Xanthoria 16
candelaria 329
elegans 329
fallax 329
parietina 12, 14
polycarpa 329
Xenokylindria 458
Xylographa
parallela 55
Zetiasplozna 439
Zoophthora 305, 313-314
15
Errata, Volume Forty-Eight
Page 542 line 24 for (1979) read (1977)
line 32 for (1977) read (1979)
Errata, Volume Forty-Nine
Page 217 line 14 for Sebacina pulverea read Exidiopsis pulverea
Publication Date for Volume Forty-Nine
MYCOTAXON for October-December 1993, 49: 1-508
was issued on December 10, 1993
516
Reviewers, Volume Fifty
The Editors express their appreciation to the following individuals who have, prior to acceptance for
publication, reviewed one or more of the papers appearing in this volume:
DS) Barr G. Guzman A. Montemartini Corte
J. M. Barrasa R. E. Halling M. Moser
A. Bellemére R. T. Hanlin M. E. Palm
G. L. Benny D. L. Hawksworth De NPegler
U. Braun P. Heinemann J. D. Rogers
O. Breuss R. A. Humber A. Y. Rossman
B. Callan B. Kendrick S. Santamaria
P. F. Cannon L. M. Kohn J. W. Sheard
V. Demoulin R. P. Korf Bb. Cy sutton
P. Diederich T. W. Kuyper C. S. Tan
N. E. El-Gholl W. F. D. Marasas I. I. Tavares
W. Gams M. Matzer T. Telleria Jorge
R. L. Gilbertson P. M. McCarthy K. H. Thorne
J. H. Ginns A. S. Methven J. M. Trappe
D. Glawe O. K. Miller D. J. Webber
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